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1997 Number 1 


Volume 51 
QL ol ISSN 0024-0966 
S41! 
589 JOURNAL 
EC NT of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


4 April 1997 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


Eric H. Merzier, President Kauri Mixxota, Vice President 

Joun M. Burns, Immediate Past Jean-Francois Lanpry, 
President Vice President 

Joun W. Brown, Vice President Davin C. Irrner, Treasurer 


Micnwack J. Smiru, Secretary 


Members at large: 

Susan J. Weller Richard L. Brown Ronald L. Rutowski 
Jon H. Shepard Charles V. Covell, Jr. Felix A. H. Sperling 
M. Alma Solis John W. Peacock Andrew D. Warren 


EpIroriAL Boarp 


Rosert K. Rossins (Chairman), Joun W. Brown (Member at large) 
Lawrence F. Gat (Journal) 
WituraM E. MILter (Memoirs) 
Puiuue J. ScHappert (News) 


Honorary LirE MEMBERS OF THE SOCIETY 


Cuares L. Remincton (1966), E. G. Munroe (1973), 
ZpravKo Lorxkovic (1980), Ian F. B. Common (1987), Joun G. FrancLemonr (1988), 
Lincoin P. Brower (1990), Doucias C. Fercuson (1990), 
Hon. Miriam Rotuscuitp (1991), CLaupe Lemaire (1992) 


The object of The Lepidopterists’ Society, which was formed in May 1947 and formally consti- 
tuted in December 1950, is “to promote the science of lepidopterology in all its branches, . . . to is- 
sue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and 
ideas by both the professional worker and the amateur in the field; to secure cooperation in all mea- 
sures” directed towards these aims. ; 

Membership in the Society is open to all persons interested in the study of Lepidoptera. All mem- 
bers receive the Journal and the News of The Lepidopterists’ Society. Prospective members should 
send to the Assistant Treasurer full dues for the current year, together with their full name, address, 
and special lepidopterological interests. In alternate years a list of members of the Society is issued, 
with addresses and special interests. 


Active members—annual dues $35.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $50.00 
Life members—single sum $1,400.00 
Institutional subscriptions—annual $50.00 


Send remittances, payable to The Lepidopterists’ Society, to: Ron Leuschner, Asst. Treasurer, 1900 
John St., Manhattan Beach, CA 90266-2608; and address changes to: Julian P. Donahue, Natural His- 
tory Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. For information about the Soci- 
ety, contact: Michael J. Smith, Secretary, 1608 Presidio Way, Roseville, CA 95661. To order back is- 
sues of the Journal, News, and Memoirs, write for availability and prices to Ron Leuschner, 
Publications Manager, 1900 John St., Manhattan Beach, CA 90266-2608. 


Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepi- 
dopterists’ Sociey, %o Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los 
Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing of- 
fices. POSTMASTER: Send address changes to The Lepidopterists’ Society, %o Natural History Mu- 
seum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. ; 


Cover illustration: “old friends’—the endemic Hawaiian nymphalid, Vanessa tameamea, feeding 
at a wilted and senescent flower of the endemic plant, Kokia kauaiensis. Original pen and ink draw- 
ing by Dale Clayton, Dept. Biol. Sciences, Southwestern. Adventist College, Keene, Texas 76059, 
USA. 


JoUuRNAL OF 
Tue LepriporTreERIstTs’ SOCIETY 


Volume 51 1997 Number 1 


Journal of the Lepidopterists’ Society 
51(1), 1997, 1-8 


PRESIDENTIAL ADDRESS 1996: ON THE BEAUTIES, USES, 
VARIATION, AND HANDLING OF GENITALIA 


JOHN M. BURNS 


Department of Entomology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, USA 


Thanks to Erynnis, I got hooked on genitalia at an impressionable age 
(Burns 1964). Although some measure of genitalic asymmetry is not rare 
in hesperiids, Erynnis is the only skipper genus with thoroughly asym- 
metric genitalia that is widespread in North America north of Mexico. I 
welcomed rampant genitalic asymmetry because it added spice to com- 
parative morphology and greatly increased the number of characters in 
structures that are taxonomically useful anyway (Figs. 1-7). 

Back when few American species of Erynnis were known and the tax- 
onomic use of genitalia was not yet in vogue, Scudder and Burgess 
(1870) seized on the asymmetric genitalia—and nothing but those geni- 
talia—in distinguishing and describing not only the known species of 
Erynnis but also a number of new ones (much to the indignant annoy- 
ance of some contemporaries who refused to accept them). The only 
valid criticism is that Scudder and Burgess did not compare enough 
genitalia to appreciate individual variation fully and so described four 
species more than once. 

The value of genitalia for distinguishing species can hardly be over- 
stated. In trying to show how to separate two large, similar looking but 
none too closely related, eastern North American species of Erynnis 
superficially, Klots (1951:pl. 29, figs. 7, 9) indicated, with photographs 
of males, the presence of two subapical white spots on the ventral hind- 
wing in E. juvenalis (Fabricius) and their absence in E. horatius (Scud- 
der & Burgess). I have examined the genitalia of his models (which 
Klots clearly labelled as such) and found both to be E. juvenalis. The 
tegumen, uncus, and dissimilar left and right valvae of E. juvenalis 
(Scudder & Burgess 1870:figs. 9, 10) depart widely from those of E. 
horatius (Scudder & Burgess 1870:figs. 13, 14). Although the uncus 
and left and right valvae of E. horatius closely resemble those of its para- 


2 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-7. Strikingly asymmetric male genitalia of the pyrgine skipper Erynnis scud- 
deri (Skinner) which ranges from southeastern Arizona, USA, to Guatemala; specimen 
from the El Tapon area, route CA 1, 5600 ft [1705 m], GUATEMALA, 8 August 1971, R. 
W. Holland (J. M. Burns genitalia no. 1360) (USNM). 1, Complete genitalia in left pos- 
terodorsolateral view. 2-4, Tegumen, uncus, and gnathos in dorsal, lateral, and ventral 
views, with posterior end to right (structures turned ninety degrees between successive 
views). 5, Right valva in right lateral view (spines included). 6, Both valvae plus aedeagus 
in dorsal view (spines omitted). 7, Left valva in left lateral view (spines included). In last 
three views, posterior end at top; aad each valva shown at two angles, ninety deste apart. 
Drawings by Robin S. Lefberg. 


VOLUME 51, NUMBER Il 3 


TABLE 1. Forewing length (mm) in genitalically identical species of Erynnis taken by 
. M. Burns in association with their larval foodplants either in (E. baptisiae) or near (E. 
lucilius) Middletown, Connecticut, USA, between 1962 and 1965. 


Species Phenotype Sex N Range Mean SE SD CV 
E. lucilius 1 3 43 12.4-15.2 13.89 OZ 0.81 5.83 
1 2 13 13.6-15.4 14.50 0.18 0.66 4.55 
2, 3 a, 13.8-15.9 15.04 0.14 0.58 3.86 
2 2 8 14.2-17.0 15.69 0.30 0.86 5.48 
E. baptisiae 1 3 o2 13.4-16.2 5s 0.13 0.72 4.76 
1 Q 3l 14.1-17.5 15.57 0.14 Om 4.95 
2 3 45 14.7-17.8 16.26 0.09 0.63 3.87 
2 2 21 16.1-18.5 Wie2s 0.15 0.70 4.06 


patric, western American sister species, E. tristis (Boisduval) (Scudder 
& Burgess 1870:fig. 15), the distal end of the tegumen, which is divided, 
forms a long, fat finger on the right side in E. horatius but a large, round 
plate on the left in E. tristis. 

Of course, the genitalia do not have to vary between species. Even in 
Erynnis, with its rich asymmetry, I have tried and failed repeatedly over 
the years to discover at least one genitalic difference between E. bap- 
tisiae (Forbes), a more southern, eastern North American differentiate 
that feeds as a larva primarily on Baptisia, but also on Lupinus, and now, 
secondarily, on an introduced Coronilla (all Fabaceae), and E. lucilius 
(Scudder & Burgess), a more northern sister differentiate that departs 
evolutionarily from its congeners by eating Aquilegia (Ranunculaceae). 
However, where these two skippers coexist in central Connecticut, I can 
statistically demonstrate a difference in the size of adults sampled in di- 
rect association with their larval foodplants, Aquilegia canadensis L. and 
Baptisia tinctoria (L.) R. Br. I know from my prior detailed analysis of 
variation in size that, within a species of Erynnis, females average larger 
than the males with which they fly and that summer generation individ- 
uals (phenotype 2) in either sex average larger than spring generation in- 
dividuals (phenotype 1) of the same sex (Burns 1964). After appropriate 
subsampling, E. baptisiae consistently averages at least one millimeter 
longer than E. lucilius in forewing length (Table 1). 

Across the genus Erynnis as a whole, I encountered such enormous 
genitalic variation that for years I implicitly accepted, or tolerated, wide 
genitalic latitude within skipper genera generally. This was a mistake. 
Genitalia are often phylogenetically constrained—so much that they of- 
fer characters of special value in grouping at the generic level (and 
above), along with those that serve in telling species apart. History did 
not help my perception, either, because pioneers (like Scudder & 
Burgess 1870, Scudder 1889, Godman & Salvin 1879-1901) in the use 
of male genitalia in distinguishing skipper species, failed to see the 


4 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


higher level information at hand and frequently put species with similar 
genitalia in different genera and species with quite different genitalia in 
the same genus. Only in the last decade have I fully realized that many 
of our much-studied and long-stable nearctic skipper genera (as well as 
many poorly studied neotropical ones) are polyphyletic and that geni- 
talia (in both sexes) provide crucial tools for sorting them out (see Burns 
1994, 1996, plus earlier papers cited therein). 

Though I lean heavily on genitalia in redefining genera, I always seek 
supporting information. After pulling three long-tailed species out of 
Polythrix and uniting them, on genitalic grounds, with tailless species in 
the distant, supposedly monotypic genus Cephise, I found that all species 
in much-expanded Cephise share a unique palpal feature and that both 
tailed and tailless species eat the same larval foodplants (Burns 1996). 

Despite its loss, “Polythrix,” a neotropical genus of similar appearing 
tailed species, is still so genitalically heterogeneous and polyphyletic that 
what remains may go in six different genera. True Polythrix are the type 
species, metallescens (Mabille), as well as kanshul Shuey and eudoxus 
(Stoll) (Burns 1996). The largest genitalically compact unit that does not 
include the type species is the asine group of six: asine (Hewitson), 
gyges Evans, hirtius (Butler), mexicanus Freeman, roma Evans, and an 
undescribed species. Another genitalically compact unit is the auginus 
group of three: auginus (Hewitson), caunus (Herrich-Schiffer), and 
an undescribed species. Three species are genitalic oddballs: ceculus 
(Herrich-Schiffer), minvanes (Williams), and octomaculata (Sepp). 

Genitalia express plenty of individual variation, which must be stud- 
ied, compared, and understood in order to interpret them correctly. As 
noted above, Scudder and Burgess (1870) were initially overimpressed 
by minor genitalic variants and described too many species of Erynnis, 
no doubt because they looked at few individuals. In the course of exam- 
ining some 12,000 genitalia during microevolutionary studies of Eryn- 
nis, I uncovered occasional major variants, the most stunning of which 
are males of E. funeralis and E. propertius (both species described as 
new by Scudder and Burgess in 1870) whose genitalia are secondarily 
symmetric: the left valva is a mirror image of the right one instead of its 
usual, highly distinctive self (Burns 1964, 1970:figs. 1-4). 

Among numerous genitalic dissections connected with an ongoing 
generic redefinition and revision, I have found—in just one of a few 
males of an undescribed neotropical species—the reverse situation 
where genitalia that are normally symmetric (except for the distal aedea- 
gus and its cornuti) are, all at once, conspicuously asymmetric in both 
the tegumen and the uncus (Figs. 8-12). Again, one of six males of an 
undescribed species in the asine group of “Polythrix” from El Salvador 
and Costa Rica has abnormal spikes directed downward from the ven- 


Fics. 8-12. Bizarre variation in male genitalia of an undescribed hesperiine skipper 
that ranges from Mexico to Colombia or Ecuador (Burns, unpubl. data). 8, 9, Normal gen- 
italia from Santa Rosa, Veracruz, MEXICO, May 1906 (J. M. Burns genitalia no. X-3037) 
(USNM). 10-12, Abnormal genitalia—with a major process (unknown in this and other, 
related, genera) arising (only on the left side) from the tegumen/uncus, above the base of 
the gnathos; a bend to the right at the distal end of the uncus; and a uniquely low dor- 
sodistal edge on the valva—from Chiriqui, PANAMA (J. M. Burns genitalia no. X-3862) 
(Museum fiir Naturkunde der Humboldt-Universitaét zu Berlin, Zoologisches Museum). 
8, 10, Tegumen, uncus, and gnathos in dorsal view. 9, 11, Complete genitalia (minus right 
valva), with vesica everted to show quadruple cornuti, in left lateral view (plus, in 9, en- 
largements of two cornuti at different angles). 12, Distal end of aedeagus, with vesica 
everted to show quadruple cornuti, in dorsal view. Drawings by Young Sohn. 


6 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


at 


Fics. 13-22. Devastating variation in male genitalia of the pyrrhopygine skipper 
Metardaris cosinga (Hewitson) from the department of Cuzco, 2850 m, PERU. 13-15, 
Normal genitalia (boiled in 10% KOH, freed from other sclerotized parts, cleaned of 
scales, muscles, etc., and stored in glycerol) in left lateral, dorsal, and ventral views; from 
5 km N Paucartambo, 13°15' S, 71°37' W, 28 August 1989, R. K. Robbins (J. M. Burns 
genitalia no. X-3044) (USNM). 16-18, Normal genitalia (dissected dry in situ) in right lat- 
eral, dorsal, and ventral views; same field data as preceding male (USNM). 19, 20, Abnor- 
mal genitalia (dissected dry in situ)—-with massive cross-fusion between the bottoms of 
the valvae, or claspers, making the genitalia totally useless—in dorsal and ventral views; 
from near Calca, 13°19' S, 72°00! W, 27 August 1989, D. J. Harvey (USNM). 21, 22, The 
whole skippers, with their dry-dissected genitalia exposed, in ventral view: normal on left, 
abnormal on right. Photographs by Carl C. Hansen. 


tral edges of its valvae near their distal ends, one on the left and two on 
the right. Taxonomists who do not set variation in a proper context 
might describe each of these two aberrant males as new for the wrong 
reasons. 

But could anyone misinterpret the male of Metardaris that leaped out 
at me from others in a batch of newly spread skippers because the distal 
end of its abdomen was strangely chafed? A victim of grossly deviant de- 
velopment that broadly joined both valvae ventrodistally (Figs. 13—22)— 
to create a sort of built-in, indestructible chastity belt—it must have 


VOLUME 51, NUMBER 1 1 


6 ¢ 


C@ 


Lb) 


Fics. 23-33. Handling, study, and storage of liberated skipper genitalia. 23, Cabinet 
drawers with rows of one-dram, screw-cap vials holding genitalia (and usually also abdom- 
inal skins) in glycerol. 24, Removing a vial from its numerical sequence in a drawer (indi- 
vidual dissection numbers on round adhesive labels on the tops of vials). 25, Two one- 
dram vials showing dissections and permanent labels with individual dissection numbers 
inside. 26, Removing dissected genitalia from a vial with forceps (here, an accessory sex- 
and-determination label accompanies the mandatory dissection label). 27, Genitalia with 
temporary tags in glycerol in a 12-depression, porcelain spotplate for critical microscopic 
study and comparison. 28, Manipulating female genitalia in a spotplate depression with 
jeweler’s forceps. 29, Directly comparing two female genitalia in a single spotplate de- 
pression. 30, Directly comparing two male genitalia in a single spotplate depression (the 
genitalia on the right [X-3044] appear, larger than life, in Figs. 13-15). 31, John Burns 
closely comparing genitalia at his work desk with variable lights and a stereomicroscope. 
32, Temporary storage of spotplates of genitalia during long, large projects: plates in a 
USNM insect drawer, with half-column pinning units upside down to serve both as dust 
covers and as surfaces for sticky notes on the dissection numbers, sexes, identities, sources, 
and peculiarities of the covered genitalia. 33, Facing pages of the requisite dissection 
notebook showing, for each genitalic preparation, a one-line entry beginning and ending 
with the dissection number, and including sex, determination, parts dissected, date of dis- 
section, time boiled in KOH, and the date, locality, and collector of the specimen, as well 
as the collection in which the specimen resides. Photographs by Chip Clark. 


spent most of its short adult life attempting basic copulatory motions 
(such as clasping with its claspers) which fizzled, ruffling it and its poste- 
rior scales. No ardent conservationist can ever censure the untimely hu- 
man capture of this specimen (Figs. 19, 20, 22) since it already had been 
naturally selected against. 

From the beginning I have handled genitalia in a novel but expedient 


8 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


manner (Figs. 23-33). I keep them free in vials because slide mounts 
may distort them and will always severely limit the angles of view and 
preclude side-by-side comparisons. I use large, one-dram screw-cap 
vials (with an inverted plastic cone in the cap) which readily hold the ab- 
dominal skin unfolded, as well as the dissected genitalia, a label with the 
dissection number, and, if desired, another with sex and determina- 
tion—all in enough glycerol to last until the next glaciation (Figs. 25, 
26). Genitalia and skins can be examined superficially within vials and 
can easily be removed from them with forceps for detailed study. Mi- 
crovials (especially older ones with cork stoppers) dry out over time, 
leak on occasion, require folding of skins and large genitalia, need to be 
pinned, may suffer stopper breakage, and are generally messier and 
harder to handle. For all-important repeated study and direct compari- 
son at each and every possible angle, I keep genitalia in glycerol in spot- 
plate depressions with temporary tags, often for years at a stretch (Figs. 
27-32). There is no better way. 


LITERATURE CITED 


BuRNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. 
Publ. Entomol. 37:1—217. 

. 1970. Secondary symmetry of asymmetric genitalia in males of Erynnis funeralis 

and E. propertius (Lepidoptera: Hesperiidae). Psyche 77:430-435. 

. 1994. Genitalia at the generic level: Atrytone restricted, Anatrytone resurrected, 

new genus Quasimellana—and yes! we have no Mellanas (Hesperiidae). J. Lepid. Soc. 

AS: Jia OoN. 

. 1996. Genitalia and the proper genus: Codatractus gets mysie and uvydixa—in a 
compact cyda group—as well as a hysterectomy, while Cephise gets part of Polythrix 
(Hesperiidae: Pyrginae). J. Lepid. Soc. 50:173—216. 

GoDMAN, F. D. & O. SALVIN. 1879-1901. Biologia Centrali-Americana; Insecta; Lepi- 
doptera-Rhopalocera. Vol. 2, 782 pp.; Vol. 3, 113 pls. 

KiorTs, A. B. 1951. A field guide to the butterflies of North America, east of the Great 
Plains. Houghton Mifflin Co., Boston. 349 pp., 40 pls. 

SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with spe- 
cial reference to New England. Publ. by the author, Cambridge, Massachusetts. Vol. 
2, pp. i-xi + 767-1774; Vol. 3, pp. i-vii + 1775-1958, 89 pls., 3 maps. 

SCUDDER, S. H. & E. BuRGEssS. 1870. On asymmetry in the appendages of hexapod in- 
sects, especially as illustrated in the lepidopterous genus Nisoniades. Proc. Boston 
Soc. Nat. Hist. 13:282—306. 


Received and accepted for publication 16 October 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 9-31 


DIVERSITY AND EVOLUTION OF TONGUE LENGTH IN 
HAWKMOTHS (SPHINGIDAE) 


WILLIAM E.. MILLER 
Department of Entomology, University of Minnesota, Saint Paul, Minnesota 55108, USA 


ABSTRACT. Hawkmoths are best known as long-tongued nectar foragers, but as 
many as one-fifth of hawkmoth species have drastically shortened tongues and do not seek 
flower nectar. Clues to tongue-length diversity and evolution have not previously been 
sought in any hawkmoth stage but the adult. Using comparative methodology, and inves- 
tigating some 150 species of New and Old World hawkmoths, I uncover correlations be- 
tween tongue length and latitude of distribution, and between tongue length and growth 
form of larval foodplants. Through north latitudes ranging from 0 to 40 or 50°, mean 
tongue length declines worldwide from more than 40 mm to 15 mm or less. Through lar- 
val foodplant growth-form indexes ranging from 2 (herbs) to 6 (trees), mean tongue- 
length similarly declines from more than 40 to less than 15 mm. I speculate for extratrop- 
ical regions that tongues have lengthened in hawkmoths that must imbibe large amounts 
of nectar as flight fuel to find inconspicuous, nonpersistent larval foodplants such as herbs, 
whereas tongues have shortened in hawkmoths that have the easier task of finding con- 
spicuous, persistent larval foodplants such as trees. Residual tongue-length variation could 
reflect miscellaneous factors operating at smaller than continental geographic scales. 


Additional key words: growth form, pollination, geographic variation, phylogenetic 
analysis. 


The long proboscis or tongue is a hallmark of Sphingidae. The tongue 
of the neotropical Amphimoeca walkeri (Bdv.), whose length can reach 
280 mm (Amsel 1938), is thought to be the longest haustellum in all In- 
secta. Also contributing to sphingid tongue lore is the story of the 
Madagascan hawkmoth Xanthopan morgani praedicta R. & J. (Kritsky 
1991). Charles Darwin observed in 1862 that the nectar of the Mada- 
gascan star orchid, Angraecum sesquipedale Thouars (Orchidaceae), is 
hidden some 290 mm deep in the blossom. A hawkmoth pollinator with 
so long a tongue was then unknown, but Darwin predicted that one 
would be found. Four decades later, Rothschild and Jordan (1903) de- 
scribed the predicted hawkmoth. According to Kritsky (1991), yet an- 
other species of Angraecum orchid with still deeper nectar has surfaced, 
and yet another hawkmoth with a still longer tongue has been pre- 
dicted! Although pollination literature often focuses on comparative 
lengths of tongues and nectar tubes, which makes one think of coevolu- 
tion, no general mechanism has been advanced to explain the diversity 
and evolution of hawkmoth tongue length. 

Hawkmoth visitors at flowers are often known by plant taxa because 
so much knowledge of nectar foraging comes from pollination studies. 
Hawkmoths insert their tongues, sometimes more of their bodies, into 
blossoms for nectar. In the process, they touch pollen-bearing anthers 
and pollen-receiving stigmas, accidentally transferring pollen within or 
among blossoms. The extensive foraging range (Linhart & Mendenhall 


10 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


1977), swift, darting flight, and nocturnal activity of many hawkmoths 
makes it more practical for an investigator to remain near flowers and 
observe arrivals, rather than follow such fleeting matchmakers to see 
what flowers they visit. Fluorescent powders, dyes, devices that enhance 
night vision, and palynological analysis of tongues and faces of captures 
have enhanced hawkmoth investigations of interest in the present study 
(Eisikowitch & Galil 1971, Kislev et al. 1972, Linhart & Mendenhall 
1977, Haber & Frankie 1989). Some of the pollinator lists by plant taxa 
that include hawkmoths are those for Angraecum spp. (Nilsson et al. 
1985), Lavandula sp. (Labiatae) (Herrera 1989), Luehea spp. (Tiliaceae) 
(Haber & Frankie 1982), Pancratium maritimum L. (Amaryllidaceae) 
(Eisikowitch & Galil 1971), and Polemoniaceae spp. (Grant & Grant 
1965). Conversely, lists by hawkmoth species of plant taxa visited or pol- 
linated (Fleming 1970, Kislev et al. 1972) are less common. 

From lists of pollinators by plant taxa, seven pollinator syndromes— 
flower types favoring different animal pollinator groups—have been 
characterized, one being the syndrome of sphingophilous or hawkmoth 
flowers (Baker & Hurd 1968, Faegri & van der Pijl 1979). Sphingo- 
philous flowers have the following traits: nocturnal anthesis or opening, 
white or pale coloration, sweet fragrance, horizontal to pendant posture, 
abundant sucrose-rich nectar, and long nectar tube (Baker & Hurd 
1968, Faegri & van der Pijl 1979, Cruden et al. 1983, Haber & Frankie 
1989). The nectar tube may be formed by various flower parts such as 
corolla, calyx, petal spur, hypanthium, or consist of a false tube formed 
by stamens and petals (Grant 1983). Hawkmoth flowers belong to the 
evolutionarily advanced stereomorphic and zygomorphic types of an- 
giosperm flowers (Leppik 1968, Crepet 1979). 

A shorter tongue than tube usually prevents nectar extraction; a 
longer tongue than tube lessens pollen removal and pollination effec- 
tiveness. Most hawkmoths use a range of available tube lengths, and 
many hawkmoth flowers are pollinated by a range of pollinator types. 
Nevertheless, comparative lengths of hawkmoth tongues and nectar 
tubes of hawkmoth flowers still interest investigators (Gregory 1963-64, 
Grant & Grant 1965, 1983a, 1983b, R. B. Miller 1978, 1981, 1985, Haber 
& Frankie 1982, 1989, Herrera 1989, Grant 1983, Martinez del Rio & 
Burquez 1986, Nilsson et al. 1985, Nilsson 1988, and others). Tube 
lengths of North American hawkmoth flowers range from nil to 175 
mm, and tongue lengths of associated hawkmoth pollinators, from 23 to 
138 mm, averaging 53 and 60 mm, respectively (Grant 1983). Corre- 
sponding statistics in a Costa Rican community are nil to 190 mm, and 
10 to 200 mm, averaging 51 and 49 mm, respectively (Haber & Frankie 
1989). The resulting tube-to-tongue ratios of 0.88 and 1.04 approximate 
those experimentally implicating current interactions as maintaining 


VOLUME 51, NUMBER 1 ll 


long tubes and long tongues (Nilsson 1988). Relations between hawk- 
moth pollinators and plants range from strong one-to-one tongue- and 
tube-length mutualisms (Nilsson et al. 1985) to more general matches 
(Leppik 1968, Proctor 1978, Feinsinger 1983, Howe 1984, Bawa 1990). 

Contrary to the popular image of long-tongued hawkmoths, several 
sphingid lineages have vestigial tongues and head musculature. Rudi- 
mentary tongues were well known by the time of Rothschild and Jordan 
(1903), and documented further by Hattich (1907), Mell (1922, 1940), 
Kernbach (1962), and Fleming (1968). Sphingidae are members of the 
suborder Glossata. One of the classical defining traits of Glossata is the 
presence of a functional proboscis (Kristensen 1984). Thus the hawk- 
moth ancestor had a functional tongue, and vestigial tongues in modern 
hawkmoths must be the result of reduction. Tongue shortening to 10 
mm or less renders hawkmoths incapable of nectar foraging (Fleming 
1968), but does not necessarily prevent them from drinking water 
(Kernbach 1962; Pittaway 1993:106). Although as many as one-fifth of 
hawkmoth species have shortened tongues and do not forage for nectar, 
interest in nonfeeding has been dwarfed by interest in nectar foraging. 
It seems unlikely that a coevolutionary hypothesis of tongue and tube 
lengths could account for tongue reduction and the order-of-magnitude 
range in tongue lengths from 2.5 mm in Laothoe juglandis (J. E. Smith) 
(Fleming 1968) to 280 mm in Amphimoeca walkeri. 

Life-system investigations of sphingids often involve either adult or 
immature stages, seldom both. One reason is that foodplants of the 
adult and immature stages usually differ. Comparison of larval and adult 
foodplant records shows that only 3 to 5 percent of hawkmoth species 
are known to use even one foodplant genus in both stages (Fleming 
1970, Hodges 1971, Grant 1983, Pittaway 1993). Clues to hawkmoth 
tongue-length diversity and evolution have never been sought in any 
but the adult stage, nor at broader than local geographic scales. 

Here I investigate tongue length on a continental spatial scale using 
large samples of both New World and Old World hawkmoth species. I 
explore correlations of tongue length with three variables: midrange lat- 
itude of hawkmoth distributions, growth-form of larval foodplants, and 
percentage of eggs that are mature at adult eclosion. I compare tongue 
length and tube length of hawkmoth flowers on a latitudinal gradient. 
Using comparative methodology, I test statistical significance of correla- 
tions of tongue length with larval-foodplant growth form, and chart the 
evolution of both traits as well as that of their correlation. 


MATERIALS AND METHODS 


In comparative studies, conventional parametric methods may inflate degrees of free- 
dom in significance testing. The reason is that traits of interest may have been inherited 
from a common ancestor rather than evolved independently by each sample taxon. This 


12 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


problem is addressed by phylogenetic nested analysis of variance and covariance (Bell 
1989, Harvey & Pagel 1991). In nested analysis, values for a given trait are nested heirar- 
chically. Here, for a trait like tongue length, populations are nested within species, spe- 
cies within genera, genera within tribes, and tribes within subfamilies. In nesting by tax- 
onomic level, species represent population ancestors, genera represent species ancestors, 
tribes represent genus ancestors, and subfamilies represent tribe ancestors. Nested 
groups are not weighted here by number of subtaxa. I use conventional parametric statis- 
tics to introduce and describe correlations, and nested analysis to estimate independent 
evolution and test statistical significance. 

Three kinds of correlation are involved in this study. First is conventional product- 
moment correlation, r estimating degree of association between a dependent variable and 
one independent variable, R when there are two independent variables. Product-moment 
correlation coefficients are usually not tested for significance here because of the degrees- 
of-freedom problem. The second kind of correlation is intrinsic correlation (Bell 1989), 
which is identical to variance-component correlation in the output of nested analysis. The 
intrinsic correlation coefficient estimates degree of association between two variables at 
various taxonomic levels, and resolves the degrees-of-freedom problem in significance 
testing. The third kind of correlation is intra- and interclass correlation. Intraclass corre- 
lation is the cumulative percentage of total variance at successively lower taxonomic or 
nesting levels. It is readily derived from variance components in the output of nested 
analysis. The intraclass correlation coefficient is used in this study to estimate indepen- 
dent evolution of a trait at a given taxonomic or nesting level, and to trace the geologic 
history of variation in the trait (Bell 1989). Finally, interclass correlation is the cumulative 
covariance or joint variation between two traits at successively lower taxonomic or nesting 
levels standardized by the geometric mean of cumulative variances for each trait (Bell 
1989). The interclass correlation coefficient is used in this study to trace the geologic his- 
tory of covariation between two traits (Bell 1989). Coefficients of product-moment corre- 
lation range strictly between 0 and 1, those of intrinsic, intra- and interclass correlation 
nominally between 0 and 1. I used the NESTED procedure of SAS (1988) to perform 
nested analyses. . 

My use of hawkmoth classification rather than phylogeny for nesting is necessary be- 
cause a cladistic or modern hawkmoth phylogeny is not available. Classifications and phy- 
logenies are not necessarily isomorphic, and nesting based on classification may obscure 
phylogenetic divergences intermediate between taxonomic levels (Harvey & Pagel 1991). 
The importance of obscured divergences, if any, will only be revealed when a modern 
phylogeny can be used in place of the classification. Hawkmoth classification has long 
been stable, which makes it a suitable surrogate for phylogeny. The classification dates 
from Rothschild and Jordan’s (1903) landmark revision of the world fauna which has not 
been appreciably altered except at the subgeneric level, a level I omit in nesting. Tribes 
Choerocampini and Macroglossini have been synonymized under the latter name (Hodges 
1971, Pittaway 1993), and I follow the resulting arrangement. For New World hawkmoths, 
nomenclature follows Hodges (1983) and D’Abrera (1986), in that order of preference. 
For Old World hawkmoths, nomenclature follows Pittaway (1993) and D’Abrera (1986), in 
that order of preference. I do not distinguish named infraspecific forms in this study ex- 
cept as populations. My geological time scale (Fig. 5) depicts taxonomic divergence in 
Sphingidae as very slow. Although inspired by Wilson (1978a, 1978b), the scaling is but a 
guess. Even if wrong in absolute time, however, it is accurate in relative time. 

Population tongue length refers to different tongue measurements for a species from 
different parts of its range. Multiple reports of tongue length are available for 41% (29/70) 
of the New World sample of hawkmoth species (Appendix 1), and for 11% (9/81) of the 
Old World sample (Appendix 2). With one exception, species tongue lengths refer to sin- 
gle reports or arithmetic means of population values including both sexes; genus tongue 
lengths refer to arithmetic means of the means of constituent species; and tribe tongue 
lengths refer to arithmetic means of the means of constituent genera. The exception con- 
cerning species tongue lengths involves the correlation between percentage of eggs that 
are mature at adult eclosion and tongue length. In this correlation, species tongue lengths 
are from females only. Sexual dimorphism in hawkmoth tongue lengths is minor, and is 


VOLUME 51, NUMBER 1 i} 


usually related to sexual dimorphism in body size. In the sample where only female tongue 
lengths are used, females average 2.0 mm longer tongues than males, ranging in individual 
species up to 5.9 mm longer (n = 18; species with tongue length >4.0 mm; Mell 1922). 

New World tongue lengths (Appendix 1) are taken from Gregory (1963-64), Fleming 
(1968, 1970), Hodges (1971), R. B. Miller (1978, 1981, 1985), Bullock and Pescador (1983), 
Grant (1983), Grant and Grant (1983a, 1983b), Martinez del Rio and Buirquez (1986), and 
Haber and Frankie (1989). Old World tongue lengths (Appendix 2) are taken from Hat- 
tich (1907), Kiinckel d’Herculais (1916), Mell (1922, 1940), Kernbach (1961), Kislev et al. 
(1972), Nilsson (1983, 1988), Herrera (1989), and Pettersson (1991). 

All published tongue lengths known to me for hawkmoths with midrange latitudes of 
0° or greater northward are included in this study, except one set from Costa Rica (Young 
1972). These appear discrepant. For example, tongue lengths reported for species of Xy- 
lophanes are about twice those for the same species elsewhere in Costa Rica and in west- 
ern Mexico (Bullock & Pescador 1983, Haber & Frankie 1989). Similar differences occur 
in Manduca, Eumorpha, Cocytius, Erinnys, Pachylia, and others. 

Tongue length takes two forms in this study: arithmetic and natural logarithmic (In). I 
use In values in the nested analyses to homogenize variance (Bullock & Pescador 1983), 
and to place differences through a wide range on one scale. Arithmetic values appear in 
scattergrams, but are plotted on logarithmic scales. Scattergram trend lines are ordinary 
least-squares fits of the exponential function y = a (10*). I add 1 to latitudes and oogenesis 
percentages in some scattergrams and analyses to avoid computational and display prob- 
lems associated with zero values. Flower-tube length also takes arithmetic form in de- 
scription, and In form in analysis. 

I use forewing length as a surrogate for body size. Live body-weight increases as the 
square of forewing length, which makes forewing length a sensitive index of body size (W. 
E. Miller 1997). Forewing lengths of sample New World hawkmoths are taken from Bul- 
lock and Pescador (1983); D’Abrera (1986), whose life-size illustrations of spread speci- 
mens I measured; and Haber and Frankie (1989). Forewing lengths of sample Old World 
hawkmoths are taken from Mell (1922) and D’Abrera (1986). In checking for body-size in- 
fluence on tongue length, I examine the correlation between tongue length and forewing 
length at the genus rather than species level. Many hawkmoth species have common an- 
cestry, which, as mentioned, may reduce the validity of significance testing at the species 
level. In checking for body-size influence on correlations between tongue length and lar- 
val foodplant growth-form, I divide sample species by forewing length into small, 
medium, and large aliquots. The respective forewing-length class limits for the New 
World sample are 17—35, 36-50, and 51-88 mm; and for the Old World, 14-29, 30-39, 
and 40-71 mm. 

Midrange latitudes serve here as comparative indexes of hawkmoth distributions. 
Midrange latitude for a species is the latitude midway on a polar axis between north and 
south extremes of the breeding distribution of combined infraspecifics, excluding erratic 
records. For a genus, midrange latitude is the arithmetic mean of midrange latitudes of 
constituent species, and for a tribe midrange latitude is the arithmetic mean of mean 
midrange latitudes of constituent genera. Midranges of New World species (Appendix 1) 
are based on Schreiber (1978). Midranges of Old World species (Appendix 2) are based 
mostly on Mell (1922) and Pittaway (1993), sometimes on D’Abrera (1986). 

Larval foodplant records for New World sample hawkmoths are from Hodges (1971), 
one source cited therein, and Janzen (1984); those for the Old World, from Mell (1922), 
Lin (1987), Pittaway (1993), and Chen (1994). Foodplants are truncated to genus, and 
non-native foodplant genera are excluded. 

Larval foodplant growth-form index (Appendices 1, 2) refers to the ae height of 
mature plants and their associated size and tendency to dominate sites. Of several avail- 
able growth-form classifications, I use a simple one similar to that in Janzen (1984), which 
recognizes five classes: tree, 25 m high; treelet, 10 to 25 m high; large-shrub, 5 to 10 m 
high; small-shrub, 3 to 5 m high; and herb (Grime 1979, Collinson 1988). Except for 
climbing foodplants, I numerically score growth forms according to Box (1981): 2 for 
herbs, 3 for small shrubs, 4 for large shrubs, 5 for treelets, and 6 for trees. Climbers are 
usually considered to have a growth-form value of zero, but here they receive values rang- 


14 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Nested analysis of variance and covariance for tongue length (In) and larval 
foodplant growth-form index. The subfamily level is omitted because of only two taxa at 
that level. Double asterisks indicate significance at P < 0.01. 


Variance 


Taxonomic Tongue Growth Intrinsic 


(nesting) level n df length form Covariance correlation 

New World 

Tribe 5 3 0.580 0.667 —0.389 —0.63 

Genus 34 29 0.474 0.707 —0.363 —0.63** 

Species 70 36 0.083 1.282 —0.148 —0.45** 

Population 124 54 0.008 0.000 0.000 0.00 

Total —— 123 1.145 2.656 —0.519 —0.39** 
Old World 

Tribe -t 2 0.467 0.420 —0.489 —1.10 

Genus 38 35 0.380 0.352 —0.082 —0.22 

Species 81 42 0.062 0.294 —0.037 —0.28 


Total a 80 0.909 2.212 —0.987 —0.70** 


ing from 2 to 4 depending on size, whether woody or herbaceous, and whether annual or 
perennial, because they may be nearly as high and large as their plant supports ( Janzen 
1975). I obtained growth-form information from standard botanical compendia: for New 
World foodplants from Fernald (1950), Croat (1978), SCS (1982), and Janzen (1984); for 
Old World foodplants from Li (1935), Tutin et al. (1964, 1968), and Keng et al. (1993). Be- 
cause many hawkmoth species use larval foodplants in several genera, foodplant growth- 
form values for species are usually means. The range of values underlying the mean 
growth-form for a hawkmoth is usually narrow, such as in Costa Rican hawkmoths ( Janzen 
1984). For a hawkmoth genus, foodplant growth-form index is the arithmetic mean of 
growth-form indexes of constituent species, and for a tribe, the arithmetic mean of 
growth-form indexes of constituent genera. 

Data concerning percentage of eggs that are mature at adult eclosion are taken from 
Mell (1922, 1940). He recorded tongue length and numbers of mature and immature eggs 
and oocytes in 4 to 46 newly eclosed females per species in Old World hawkmoths (Ap- 
pendix 2). He verbally described the resulting relation. I elaborate his observations with 
product-moment correlation analysis at species, genus, and tribe levels. 


RESULTS 


The assembled New World tongue-length sample consists of 124 ob- 
servations on 70 species in 34 genera, 5 tribes, and 2 subfamilies, with 
multiple observations on 24 species (Table 1, Appendix 1). Although 
nesting extended to subfamily, no subfamily results are given for any 
variable because of only two taxa at that level. Tongue length in the 
New World sample varies inversely with latitude of midrange distribu- 
tion at species, genus, and tribe levels, with product-moment correla- 
tion coefficients ranging between —0.44 and —0.57 (Fig. 1). Although 
these correlation coefficients were not tested for significance, they are 
judged to reflect a real relation because of similar signs and values at the 
different taxonomic levels. At the species level, mean tongue lengths at 
north latitudes of 0° (northern Brazil), 20° (southern Mexico), and 40° 
(central U. S.) are 50, 25, and 15 mm, respectively. 


VOLUME 51, NUMBER 1 15 


Tongue length (T) 
(mm) 


1000 


T = 43.6 x 10 (-9-011 L) 
Species r=-0.57 


Growth-form index < 3 


1000 


100 


1000 


PSGelss4 ly ON 
r=-0.44 


100 


) 10 20 30 40 50 
Midrange latitude (degrees N + 1) (L) 


Fic. 1. Relation of tongue length to midrange latitude in New World hawkmoths at 
species, genus, and tribe levels. Points are means except where only one tongue-length 
value is available. The species equation describes the center trend line and encompasses 
all species points. Significance values are omitted because of problematic degrees of free- 
dom as explained in text. Dotted lines at 10 mm represent length below which tongues 
are believed to be nonfunctional. 


16 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Tongue length (T) 
(mm) 


1000 


: T = 33.6 x 10 (~ 9.009 L) 
Species =- 0.28 


Growth-form index <3 
\ 


1000 


T = 43.8 x 10 (-0.015L) 
=- 0,37 


100 


10 


1000 
T = 2275.0 x 10 (- 9.078 L) 


r=-0.98 


100 


10 


0 10 20 30 40 50 
Midrange latitude (degrees N) (L) 


Fic. 2. Relation of tongue length to midrange latitude in Old World hawkmoths at 
species, genus, and tribe levels. Points are means except where only one tongue-length 
value is available. The species equation describes the center trend line and encompasses 
all species points. Significance values are omitted because of problematic degrees of free- 
dom as explained in text. Dotted lines at 10 mm represent length below which tongues 
are believed to be nonfunctional. 


VOLUME 51, NUMBER 1 iy 


The assembled Old World tongue-length sample consists of 92 obser- 
vations on 81 species in 38 genera, 4 tribes, and 2 subfamilies, with mul- 
tiple observations on 9 species (Table 1, Appendix 2). Although nesting 
extended to subfamily, no subfamily results are given for any variable 
because of only two taxa at that level. Old World tongue lengths also 
vary inversely with latitude of midrange distribution at species, genus, 
and tribe levels, with product-moment correlation coefficients ranging 
between —0.28 and —0.98 (Fig. 2). At the species level, mean tongue 
lengths at north latitudes of 0° (Borneo), 25° (southern China), and 50° 
(central Eurasia) are 42, 26, and 12 mm, respectively. The correlation 
here is also judged to reflect a real relation for the same reasons given 
above for the New World sample. New and Old World sample species 
represent 14% (151/1050) of the world hawkmoth fauna (D’Abrera 1986). 

In both New and Old World tongue-length correlations with latitude, 
the trend lines for larval foodplant growth-form indexes <3 and >5 re- 
veal a tendency for tongue lengths in these narrow index ranges to 
shorten as latitude increases (Figs. 1, 2). Thus the effect of latitude 
seems to operate regardless of foodplant growth-form. 

Unlike tongue length, nectar-tube length appears to remain constant 
rather than shorten with increasing north latitude. In a community of 
hawkmoth flowers in Costa Rica (about 10° north latitude), mean tube 
length is 50 mm (n = 30; Haber & Frankie 1989), and in all known U. S. 
hawkmoth flowers (centering at 40° north latitude), mean tube length is 
55 mm (n = 124; Grant 1983). The difference, 5 mm, is not significant 
(P = 0.23, Student t-test of difference in tube lengths [In]). 

Tongue length is inversely correlated also with larval foodplant 
growth-form index at species, genus, and tribe levels. In the New World 
sample, the product-moment correlation coefficients range between 
—0.43 and —0.51 (Fig. 3). At the species level, mean tongue lengths at 
foodplant growth-form indexes of 2 (herbs), 4 (large shrubs), and 6 
(trees) are 53, 27, and 14 mm, respectively. In the Old World sample, 
product-moment correlation coefficients range between —0.59 and 
—0.84 (Fig. 4). At the species level, tongue lengths at foodplant growth- 
form indexes of 2, 4, and 6 are 41, 17, and 8 mm, respectively. As be- 
fore, these correlation coefficients were not tested for significance, but 
they are judged to reflect a real relation because of similar signs and val- 
ues at the different taxonomic levels. The mean tongue lengths for a 
given growth-form index vary some between New and Old World sam- 
ples, but the relations are similar in form. 

Previous authors report significant correlations between tongue 
length and body size in local hawkmoth assemblages (Bullock & 
Pescador 1983, Haber & Frankie 1989). Investigation here reveals that 
positive correlations between tongue length (T) and body size (F) are 


Tongue length (T) 


(mm) 
1000 
T = 102.5 x 10 (9-144 I) 
Species Sainic 
page oder a Medium-bodied _ 
100 Sc L Small-bodied 
oO Bo ! 
oO Ajo U2} 
Lo fo 


Dig 


10 


1000 3 
= 86:2)x 10150 am) 
r=-0.43 


100 


10 


1000 


T = 128.9 x 10 (9-163 !) | 
r=-0.51 


100 | 


10 


2 3 4 5 6 
Larval foodplant growth-form index (|) 


Fic. 3. Relation of tongue length to larval foodplant growth-form in New World 
hawkmoths at species, genus, and tribe levels. Points are means except where only one 
tongue-length value is available. The species equation represents all body sizes. Small, 
medium, and large body-size classes are defined, respectively, by forewing lengths of 
17-31 mm (open circles), 32-46 mm (triangles), and 50-88 mm (open squares). Respec- 
tive equations and product-moment correlation coefficients are T = 45.0 x 10°23! 
(xr = —0.45), T = 86.0 x 10-°.!28! (ry = —0.51), and T = 152.2 x 10-9171 (¢ = —0.51). Signifi- 
cance values are omitted because of problematic degrees of freedom as explained in text. 
Dotted lines at 10 mm represent length below which tongues are believed to be nonfunc- 
tional. 


Tongue length (T) 


(mm) 
1000 > 
: T = 94.6 x 10 (9-181 1) 
Species r=-0.59 
Large-bodied Medium-bodied 
100 & 1 a 


oO Small-bodied 


! 
| o,4 
oO 


10 


1000 
TS Teri 2919 COI 
=- 0.66 


100 


10 


100 T = 270.5x 10-2781) 
r=-0.84 
100 
10 
1 


2 3 4 5 6 
Larval foodplant growth-form index (I) 


Fic. 4. Relation of tongue length to larval foodplant growth-form in Old World hawk- 
moths at species, genus, and tribe levels. Points are means except where only one tongue- 
length value is available. The species equation represents all body sizes. Small, medium, 
and large body-size classes are defined, respectively, by forewing lengths of 14-29 mm 
(open circles), 30-39 mm (triangles), and 40-71 mm (open squares). Respective equa- 
tions and product-moment correlation coefficients are T = 79.8 x 10 ~°!8¢! Ges O07), 40 = 
165.0 x 10 -0.2581 (r = —0.76), and T = 69.1 x 10 ~9."12I (r = —0.34). Significance values are 
omitted because of problematic degrees of freedom as explained in text. Dotted lines at 
10 mm represent length below which tongues are believed to be nonfunctional. 


20 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


pervasive both above the species level and at broader geographic scales. 
For genera, the New World sample yields T = 4.52 x 109-018F (r = 0.61, 
n = 34); the Old World sample, T = 7.53 x 10°°UF (7 = 0.32, n = 38). If 
the degrees-of-freedom problem is assumed to be small at the genus 
level, then the significance values for these product-moment correla- 
tions are P < 0.01 and 0.05, respectively. 

Despite the influence of body size on tongue length, the tongue- 
length correlations with foodplant growth-form are not merely an out- 
come of different hawkmoth body sizes. Tongue length decreases in all 
body-size ranges as foodplant growth-form index increases. In the three 
body-size aliquots of New World species, product-moment correlations 
range between —0.49 and —0.58, compared with —0.48 for all New World 
species (Fig. 3); in the three body-size aliquots of Old World species, 
they range between —0.34 and —0.78, compared with —0.59 for all Old 
World species (Fig. 4). The most telling evidence that growth form op- 
erates in all body-size ranges is that regression slopes in all six New and 
Old World aliquots are negative. Multiple regression analysis echoes 
this conclusion in its standardized slope coefficients, which reveal the 
relative influence of independent variables (SYSTAT 1992). These coef- 
ficients for foodplant growth-form and body size in New World species 
are —0.44 and 0.61, respectively (r = 0.77); and in Old World species 
—0.65 and 0.32, respectively, (r = 0.67). 

Similarity of tongue-length change with latitude and larval foodplant 
growth-form index at three taxonomic levels (Figs. 1-4) suggests not 
only real relations, but relations with a long history. Both suggestions 
are confirmed for growth-form index by nested analyses (Table 1, Fig. 
5), and for latitude by extension and inference. Overall intrinsic corre- 
lations between tongue length and growth-form index for New and Old 
World hawkmoth samples are —0.39 and —0.70, respectively (P < 0.001) 
(Table 1). Similar intrinsic correlations appear at the tribe level, which 
represents truly ancient ancestors. The respective New and Old World 
tribe covariances of —0.389 and —0.489 are the highest of any taxonomic 
level (Table 1). The histories of variation in tongue length and foodplant 
growth-from, as well as that of their covariation, show little change since 
genera diverged perhaps 15 million years before present (Fig. 5). De- 
spite tongue-length relations with latitude and larval foodplant growth- 
form, it must be acknowledged that significant tongue-length variation 
remains unexplained (Figs. 1-4, Table 1). 

The sample concerning percentage of eggs that are mature at adult 
eclosion in relation to tongue length consists of 26 Old World species in 
14 genera, 2 tribes, and 1 subfamily (Appendix 2). Mell (1940) con- 
cluded from this sample that oogenesis at adult eclosion is more ad- 
vanced the shorter the tongue. Mell’s conclusion also applies at the 


VOLUME 51, NUMBER 1 21 


Correlation 


Qua- 
0.2 fer 


Oligocene Miocene Pliocene nary 

0.0 

1.0 

0.8 

0.6 Species 

0.4 

Tribe 
Qua- 

0.2 L ter- 
Oligocene Miocene Pliocene _—inary 

0.0 

30 20 10 0 


Million years before present 


Fic. 5. Evolution of tongue length (solid circles, dashed lines), larval-foodplant 
growth-form (open circles, dotted lines), and their covariation (triangles, solid lines) at 
different taxonomic levels. Points for tongue length and foodplant growth-form represent 
intraclass correlations, and points for covariation represent interclass correlations, as ex- 
plained in text. The geologic time scale is a best guess based on Wilson (1978a, lo), 


genus level, and, as far as the data go, at the tribe level (Fig. 6). These 
results point to another ancient relation. At the species level, mean per- 
centage of eggs mature at eclosion for tongue lengths of 20, 50, and 80 
mm, are 30, 15, and 7, respectively. If total egg production were known 
and incorporated, it would probably intensify the relation. That is, in 
long-tongued individuals, oocytes undetected at adult eclosion, or 
formed afterwards, would likely grow and mature from resources 
gained by nectar foraging. The link between tongue length and repro- 
ductive readiness at adult eclosion confirms that tongue length is inti- 
mately involved in hawkmoth life-system evolution. 


22 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Eggs mature 
at eclosion 
(% + 1) (E) 


100 


10 : 
Species 
Suns /estiexeni ine COM) 
: r=-0.48 E = 54.9 x 19 00127) 
: “r=-0.69 
{ : ° 
0 20 40 60 80 100 


Mean female tongue length (mm) (T) 


Fic. 6. Relation of egg maturity at eclosion to tongue length in Old World hawk- 
moths. Points are means for females. Closed circles represent species, open circles gen- 
era, and open squares tribes. Data from Mell (1922, 1940). Dotted line at 10 mm a 
sents length below which tongues are believed to be nonfunctional. 


DISCUSSION 


A hypothesis of hawkmoth tongue-length diversity and evolution 
must account for both tongue lengthening and shortening as well as for 
the new findings in this study. I summarize these findings as follows. 
Mean tongue length decreases with increasing north latitude worldwide 
(Figs. 1, 2), whereas tube length of hawkmoth flowers appears to remain 
constant. Mean tongue length decreases with increasing larval food- 
plant growth-form index worldwide (Figs. 3, 4, Table 1). Further, the 
shorter the tongue, the greater the percentage of eggs that are mature 
at adult eclosion (Fig. 6). These relations are ancient, their trends hav- 
ing formed before the divergence of genera, postulated as occurring in 
middle Miocene time, some 15 million years before present (Fig. 5). 

At least three-quarters of the 25 tree genera used as larval foodplants 
by hawkmoths in this study are recorded as fossils from the middle 
Miocene or earlier (Leopold & MacGinitie 1972, Tanai 1972, Vakh- 
rameev 1991). Also, plants with stereomorphic flowers, or still more ad- 
vanced zygomorphic flowers with long nectar tubes, existed by the mid- 
dle Miocene also (Leppik 1968, Proctor 1978, Crepet 1979). Thus the 


VOLUME 51, NUMBER 1 25 


No. phanerogam 
species locally (N) 


N = 9639 - 128 L 
= - 0.94 


0 20 40 60 80 
Latitude (degrees N) (L) 


Fic. 7. Species richness of local seed-plant floras in the northern hemisphere as re- 
lated to latitude. Data from Rejmanek (1976). 


potential for larval foodplants of high growth-form index, and adult 
foodplants with long-tubed flowers, to influence tongue length seems 
easily coextensive with the history of tongue-length variation on the 
postulated geologic time scale (Fig. 5). The latitudinal gradient in 
tongue length is equally or more ancient; the position of the continents 
relative to the equator has not changed greatly since the Paleocene, 
some 60 million years before present (Smith & Briden 1977). 

No doubt many plausible hypotheses of tongue length diversity and 
evolution could be given. I favor an admittedly anthropocentric possi- 
bility, which focuses on larval foodplant finding, a process that remains 
to be studied. Especially in extratropical landscapes, herb foodplants 
(growth-form index 2) may be harder to find than tree foodplants 
(growth-form index 6). Moreover, foodplants of any growth-form index 
may be easier to find at higher north latitudes today because there is lit- 
tle doubt that patch size increases with increasing north latitude. This 
increase in patch size is as yet poorly quantified, and can best be visual- 
ized as a function of the polar-equatorial gradient of species richness in 
plants (Fig. 7). In this gradient, the number of seedplant species in local 
floras decreases at higher latitudes worldwide (Rejmanek 1976, Currie 
& Paquin 1987). As floras diminish in species richness toward polar 
regions—from near 10,000 species at the equator to one-fifth that num- 


24 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ber in central North America and central Eurasia (Fig. 7)—woody 
plants, and probably vascular plants in general, may occur in larger 
stands or patches (Dobzhansky 1950, Bourgeron 1983, Longman & 
Jenik 1987). Tropical forests often contain 60 to 1000 tree species per 
ha. In such forests, it seems physically impossible for very many indi- 
viduals of one kind of tree to occur contiguously. Although unclear 
whether patch size of all plant growth-forms in the tropics would be 
similarly affected, small patch size for trees seems likely to prevail. In 
contrast, temperate forests often have only one or a few tree species per 
ha, rarely more than 10, and large patch sizes for trees and other plant 
growth-forms prevail. 

Most hawkmoth adults are heavy bodied, and their energy expendi- 
ture in flight is enormous, with hovering consuming about | mg of sugar 
eg! body weight min“ (Heinrich 1983). For hawkmoths that must find 
nonpersistent, inconspicuous foodplants of low growth form in small 
patches (Figs. 2, 4, 7), and whose eggs are mostly immature at eclosion 
(Fig. 6), long tongues and nectar foraging are essential. The longer the 
tongue, the greater and faster the access to nectar in flowers of different 
depths, and the greater the chances of mutualistic specialization (Nils- 
son 1988, Haber & Frankie 1989, Herrera 1989). Also, the deeper the 
nectar, the more of it plants produce (Haber & Frankie 1989). In con- 
trast, long tongues and nectar foraging may be superfluous for hawk- 
moths that have the easy task of finding persistent, conspicuous larval 
foodplants of high growth form in large patches (Figs. 2, 4, 7), and 
whose eggs are mostly mature at eclosion (Fig. 6). 

The association of tongue shortening and larval feeding on trees in 
Glossata is not unique to hawkmoths (W. E. Miller 1996). Larvae of Ly- 
mantriidae and Saturniidae feed almost exclusively on trees or other 
woody plants, and their adults do not feed (Ferguson 1971—72, 1978, 
Janzen 1984, Schaefer 1989, Stone 1991). The same is true for many 
subgroups in other families such as Geometridae. Another trait associ- 
ated with larval tree-feeding and loss of adult feeding capability in 
some Glossata is reduction in female flight capability. Flightless females 
dramatically demonstrate that flight is not essential when larval food- 
plants are trees occurring in large stands or patches (Gohrbandt 1940, 
Barbosa et al. 1989, Sattler 1991). 

Although the foregoing speculative hypothesis involves mechanisms 
operating at continental geographic scales, it does not rule out other 
mechanisms of tongue-length adjustment operating at local scales. 


ACKNOWLEDGMENTS 


I thank L. S. Fink, R. C. Lederhouse, and R. K. Robbins for useful comments and dis- 
cussion on an oral presentation of the main ideas in this paper; also D. H. Janzen, S. J. 
Weller, and an anonymous reader for challenging manuscript reviews. 


VOLUME 51, NUMBER 1 95 


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Received for publication 9 June 1995; revised and accepted 7 January 1996. 


APPENDIX 1. Taxa and variable values for sample New World hawkmoths. Sources are 
given in the Methods section. For mean tongue length, values without decimals are single 
observations in a single report; values with decimals are means of multiple observations 
at one location; values with decimals and SD are means of observations at multiple loca- 
tions enumerated by (n). 


Mean tongue Midrange Mean larval 
length + SD latitude foodplant growth 
Species (mm) (°N) form index 

Sphinginae: Sphingini 

Agrius cingulata (F.) 99.2 + 4.6 (4) 0) 2.0 
Cocytius antaeus (Drury) 139 20 5.0 
Ceratomia amyntor (Gey.) 12.0 38 6.0 
C. catalpae (Bdv.) 4.4 36 ; 6.0 
C. undulosa (WIlk.) 9.8 40 6.0 
Dolba hyloeus (Drury) 32 38 4.3 
Lapara bombycoides Wk. S)) 42 6.0 
Manduca barnesi (Clark) 52.0 15 4.5 
M. corallina (Drc.) 56.8 + 1.6 (2) 15 5.5 
M. dilucida (Hy. Edw.) 43.2 + 0.4 (2) 13 5.0 
M. florestan (Cram.) 72.0 + 15.6 (3) 5 3.1 
M. lefeburei (Guér.) 51.8 + 1.2 (2) 0. 5.0 
M. muscosa (R. & J.) 86.0 25 DD 
M. quinquemaculata (Haw.) 110.0 + 14.9 (4) 40 2.0 
M. occulta (R. & J.) 68.0 11 22, 
M. rustica (F.) 138.3 + 5.1 (3) 0 3.0 
M. sexta (L.) 89.0 + 6.4 (A) 33 2.0 
Neococytius cluentius (Cram.) 228.5 + 37.5 (2) ili 3.0 
Sphinx chersis (Hbn.) 50.4 + 7.6 (5) 40 a 
S. drupiferarum J. E. Sm. 52.0 + 11.3 (2) 4] 5.5 
S. eremitoides Stkr. 39.0 40 2.0 
S. kalmiae J. E. Sm. 40 39 5.2 
S. libocedrus Hy. Edw. 45 30 4.0 
S. sequoiae Bdv. 23.0 35 6.0 
S. vashti Stkr. 60.0 + 2.6 (3) 40 : 3.0 


Sphinginae: Smerinthini 


Laothoe juglandis (J. E. Sm.) 2.5 40 6.0 
Pachysphinx modesta (Harr.) 4.0 Al 5.5 


VOLUME 51, NUMBER 1 


APPENDIX I. 


Species 


Paonias excaecatus (J. E. Sm.) 
P. myops (J. E. Sm.) 
Protambulyx strigilis (L.) 
Smerinthus cerisyi Kirby 

S. jamaicensis (Drury) 


Macroglossinae: Dilophonotini 


Aellopos clavipes (R. & J.) 

A. fadus (Cram.) 

A. titan (Cram.) 

Callionima falcifera (Gehl.) 
Erinnyis alope (Drury) 

E. ello (L.) 

E. lassauxii (Bdv.) 

E. obscura (F.) 
Eupyrrhoglossum sagra (Poey) 
Hemaris diffinis (Bdv.) 

H. thysbe (F.) 

Isognathus rimosus (Grt.) 
Nyceryx coffeae (WIk.) 
Pachylia ficus (L.) 

P. syces (Hbn.) 

Pachylioides resumens (WIlk.) 
Perigonia lusca (F.) 

Phryxus caicus (Cram.) 
Pseudosphinx tetrio (L.) 


Macroglossinae: Phillampelini 


Eumorpha achemon (Drury) 
E. anchemola (Cram.) 

E. fasciata (Sulz.) 

E. labruscae (L.) 

E. pandorus (Hbn.) 

E. vitis (L.) 


Macroglossinae: Macroglossini 


Amphion floridensis Clark 
Cautethia spuria (Bdv.) 

C. yucatana Clark 

Darapsa myron (Cram.) 

D. pholus (Cram.) 

Deidamia inscripta (Harr.) 
Hyles lineata (L.) 
Proserpinus terlooii Hy. Edw. 
Sphecodina abbottii (Swain.) 
Xylophanes pluto (F.) 

X. porcus (Hbn.) 

X. tersa (L.) 

X. turbata (Hy. Edw.) 


Continued. 


Mean tongue 
length + SD 


(mm) 


12.2 


14.8 


23.0 + 1.4 (2) 


Midrange 
latitude 
(°N) 


jp 


29 


Mean larval 
foodplant growth 


form index 


30 


APPENDIX 2. Taxa and variable values for sample Old World hawkmoths. Sources are 
given in the Methods section. For mean tongue length, values with decimals are means of 
multiple observations from one location; values with decimals and SD’s are means of ob- 
servations from multiple locations enumerated by (n). Superscript b = species used here 
in the correlation between percentage of eggs that are mature at adult eclosion and tongue 
length (Fig. 6). Superscript c = tongue so short as to be nonfunctional according to Mell 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(1922, 1940); mean of all nonfunctional tongues measured by him was assigned. 


Species 
Sphinginae: Sphingini 


Acherontia atropos (L.) 
A. lachesis (F.)> 

A. styx (Westw.)> 

Agrius convolvuli (L.)> 
Dolbina inexacta (WIlk.)> 
Meganoton analis (Fldr.)> 
M. rufescens Btlr.> 
Psilogramma increta (Wlk.)> 
P. menephron (Cram. )> 
Sphinx calligineus Btlr.> 
S. ligustri L. 

S. pinastri L. 


Sphinginae: Smerinthini 


Ambulyx kuangtungensis (Mell) 
A. liturata Btlr.> 

A. ochracea Btlr.> 

A. schauffelbergeri B. & G.» 

A. sericeippenis Btlr.> 

A. subocellata F\dr.> 
Amplypterus panopus (Cram. )> 
Clanis bilineata (Wlk.)> 

C. undulosa Moore? 

Cypa decolor Wlkr.» 

Laothoe populi (L.) 
Leucophlebia lineata Westw. 
Marumba cristata (Btlr.)> 

M. dyras (Wlk.)> 

M. gaschkewitschi (B. & G.)> 
M. spectabilis (Btlr.)> 

Mimas tiliae (L.) 

Parum colligata (Wlk.)> 
Polyptychus trilineatus Moore? 
Smerinthulus chinensis R. & J.» 
S. pallidus Mell 

Smerinthus ocellatus (L.) 

S. planus Wk.» 


Macroglossinae: Dilophonotini 


Cephonodes hylas (L.) 
Hemaris staudingeri (Leech) 
Sataspes infernalis (Westw.) 
S. tagalica Bdv. 


Mean tongue 
length + SD 


(mm) 


28.5 + 1.4 (2) 


Midrange 
latitude 


(°N) 


Mean larval 
foodplant growth 


form index 


VOLUME 51, NUMBER Il 


APPENDIX 2. 


Species 


Macroglossinae: Macroglossini 


Acosmerycoides leucocraspis (Hamp. ) 
Acosmeryx castanea R. & J. 

A. naga (Moore) 

A. pseudomissa Mell 

A. sericeus (Wlk.) 

Ampelophaga rubiginosa B. & G. 
Aspledon himachala (Btlr.) 

A. hyas (WIk.) 

Cechenena lineosa (WIlk.) 

C. minor (Btlr.) 

Daphnis hypothous (Cram.) 
Deilephila elpenor (L.) 

D. porcellus (L.) 

Hayesiana triopus (Westw.) 
Hippotion boerhaviae (F.) 

H. rafflesi (Btlr.) 

Hyles gallii (Rtmbg.) 

H. livornica (Esper) 
Macroglossum bombylans (Bdv.) 
M. corythus Wk. 

M. passalus (Drury) 

M. pyrrhostictum (Btlr.) 

M. sitiene (WIlk.) 

M. stellatarum (L.) 

M. troglodytus (Bdv.) 
Micracosmeryx macroglossoides Mell 
Panacra busiris Wk. 

P. mydon Wik. 

Pergesa actea (Cram.) 

Rhagastis albomarginatus (Roths.) 
R. mongoliana (Btlr.) 

R. olivaceae (Moore) 
Sphingonaepiopsis pumilio (Bdv.) 
Sphecodina caudata (B. & G.) 
Theretra alecto (L.) 

T. clotho (Drury) 

T. japonica (Orza) 

T. latreillei (MacLeay) 

T. nessus (Drury) 

T. oldenlandiae (F.) 

T. pallicosta (Wlk.) 

T. silhetensis (Wlk.) 

T. suffusa WIk. 


Continued. 


Mean tongue 


length + SD 


27.9 
30.4 
32.0 
27.9 
30.2 
28.2 
15.4 
14.6 
50.3 
44.2 
43.5 
DNS) 
18.5 
33.0 
31.4 
37.0 
25.4 
24.4 
28.0 
33.3 
32.3 
31.4 
31.8 
26.4 
26.9 
17.2 
38.4 
32.0 
73.5 
29:2 
24.0 
49.5 
12.0 
18.9 
52.7 
18.5 
Die 
40.2 
59.5 
32.8 
37.2 
30.5 
53.9 


(mm) 


+ 0.6 (3) 
+ 0.3 (2) 


+ 0.6 (3) 


Midrange 


latitude 
(°N) 


31 


Mean larval 
foodplant growth 


form index 


Journal of the Lepidopterists’ Society 
51(1), 1997, 32-46 


THE IDENTITY OF FILATIMA ORNATIFIMBRIELLA 
(CLEMENS 1864) (GELECHIOIDEA: GELECHIIDAE) 


RONALD W. HODGES 
AND 


D. ADAMSKI 


Systematic Entomology Laboratory, Agricultural Research Service, USDA, c/o U.S. 
National Museum of Natural History, Washington, D.C. 20560, USA 


ABSTRACT. Adults of Filatima ornatifimbriella (Clemens 1864) can be confused 
with three other species, two of which, Filatima occidua and Filatima adamsi are new. 
Filatima ornatifimbriella (Clemens) and F. xanthuris (Meyrick) are redescribed, and com- 
plete synonymy for each species is given. A lectotype is designated for Gelechia amor- 
phaeella Chambers 1877. Photographs of wing patterns and scanning electron micro- 
graphs of diagnostic wing features are included. A key to the species is provided in 
conjunction with illustrations of male and female genitalia. 


Additional key words: Lepidoptera, Gelechiidae, Filatima, ornatifimbriella. 


Classifications based on single character systems often result in recog- 
nition of polyphyletic groups. More natural classifications at the species 
and generic levels for Lepidoptera result when genital characters are 
analyzed together with head and venational characters. This realization 
by previous lepidopterists enabled them to recognize Gelechia Hiibner 
as a composite taxon and prompted a transfer of species from this con- 
cept to existing or newly recognized genera (Busck 1939, Sattler, 1960). 
Filatima (Busck 1939) was recognized as a result of these observations. 

Filatima is characterized as follows: labial palpus recurved to near 
vertex, third segment nearly as long as second segment, anteroventral 
scales of second segment divergent, forming a “furrowed brush;” ocel- 
lus present; forewing with M,, M,, and CuA, somewhat approximate; 
hindwing with M, and CuA, connate; Rs and M, approximate; male 
hindwing often with curtain scaling (Busck 1939 [”curtain scales”]; 
Clarke 1947 [curtain scaling]) within area from wing base to slightly be- 
yond end of cell and between Sc+R, and Rs (Figs. 8-9), other sex scales 
on posterior half of discal cell and basal portion of cells M,—-CuA, and 
CuA,—CuA,, and part of anal area (Figs. 8-10); male genital capsule en- 
closed within eighth segment; uncus hood shaped; gnathos narrow and 
somewhat recurved; costal lobe of valva elongate and narrow; saccular 
lobe of each valva asymmetric; vinculum rounded; aedeagus with lateral 
sclerite from zone and several other internal sclerites; ventral surface of 
eighth tergum with paired, basolateral scale pencils, dorsal surface usu- 
ally with prominent, long, anteriorly directed scales arising posterome- 
dially; female genitalia with antrum sclerotized or membranous; incep- 
tion of ductus seminalis on anterior part of accessory bursa; posterior 


VOLUME 51, NUMBER 1 33 


part of corpus bursae and accessory bursa often with dense micro- 
trichia; signum present or absent. 

Most Filatima are Holarctic in distribution with the greatest species’ 
diversity in semiarid areas of western United States and Mexico. Their 
larvae are leaf tiers on Acacia Mill., Amorpha L., Astragalus L., Cercid- 
ium Tul., Glycyrrhiza L., Leucaena Benth., Lupinus L., Mimosa L., 
Prosopis L., Robinia L., Thermopsis Robt. Brown, Vicia L. (Fabaceae); 
Prunus L., Purshia DC. (Rosaceae); Ribes L. (Saxifragaceae); Vaccinium 
L. (Ericaceae); Phoradendron Nutt. (Loranthaceae); Salix L. (Salica- 
ceae); and Betula L. (Betulaceae) (host information taken from speci- 
men label data on material in USNM collection). 

The closely similar wing patterns of Filatima ornatifimbriella 
(Clemens) and F. xanthuris (Meyrick) have led to their being misidenti- 
fied in museum collections as well as confused with the other species 
described herein. The goals of this study are to clarify the taxonomic re- 
lationships among these species and to provide efficient means for their 
identification. 


METHODS 


The Methuen Handbook of Colour (Kornerup & Wanscher 1978) was 
used as a color standard for the description of the adult vestiture. Geni- 
talia were dissected as described by Clarke (1941), except mer- 
curochrome and Chlorazol Black E (Kodak) were used as stains. In ad- 
dition, the ventral part of the genital capsule was separated from the 
dorsal part so both aspects could be examined with minimal distortion 
and confusion related to overlap (Fig. 15 contrasted with Figs. 16-18), 
following Pitkin (1984) and Huemer (1987). Terminology of genitalia 
follows Klots (1970). Pinned specimens and genital preparations were 
examined with stereoscopic and compound microscopes. Wing mea- 
surements were made using a hand-held micrometer. Specimens for 
SEM studies were mounted on stubs using double-sticky tape and 
coated with gold-palladium for five minutes with a HUMMER-X sput- 
ter coater. Wing scale ultrastructure was studied using an AMRAY 1810 
Scanning Electron Microscope. 


RESULTS 
Key to the Species of Filatima Confused with F. ornatifimbriella 


A ac NN i dans SE a ile ca a MIN cai ioe als ag hn Ryd aiedensle cwea leer mtagdgae neils Sar Slintwls 2 

Paemnalle . 5 5 fo Pe Ee ee et cy oe oP ae eo) Pee ee een ene Ce eee 4 

2. Valval saccular lobes slightly asymmetrical (Fig. 16) ............... ornatifimbriella 

- Valval saccular lobes greatly asymmetrical (Figs. 17-18) ...................4. 3 
3. Lateral sclerite from zone projected from aedeagus, curved, ventral sclerite 

Savellipticalywithoutajsenateredge (Mig; 20) a.) gece 32 tee ee xanthuris 


- Lateral sclerite from zone not projected from aedeagus, straight; ventral sclerite 
Sutera with ,asexrate|edge (Mig. 20) i. sinus. 8 2 oe sue wie eee adamsi 


34 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


4: Antrum!sclerotized,/signum present (Migs, 22,.24)) 03.2 ee 5 
- Antrum membranous jsignum/ absent (Hig 23)) > 5225 44.) eee xanthuris 
5. Membrane adjacent to anterior apophyses forming a deep invaginated pocket, 
microtrichiatwithiml(hig22)0> 2a ane ornatifimbriella 
- Membrane adjacent to anterior apophyses not forming an invaginated pocket, 
without microtrichia. (Fig. 24) |..." syn. . 1.) eee ee 8 occidua 


Filatima ornatifimbriella (Clemens 1864) 
(Figs. 1, 11-12, 16, 19, 22) 


Gelechia ornatifimbriella; Clemens 1864:420; Chambers 1878b:145; Smith 1891:102; Dyar 
1903:517; Busck 1903:899; Barnes & McDunnough 1917:157; Forbes 1923:271; Meyrick 
1925:84; McDunnough 1939:71. 

Filatima ornatifumbriella; Hodges 1983:23. 

Gelechia unctulella; Zeller 1873:257-8; Chambers 1878b:147; Smith 1891:102; Dyar 
1903:513; Busck 1903:878; Barnes & McDunnough 1917:157 [jr. syn. of ornatifim- 
briella|; Forbes 1923:267; Meyrick 1925:84 [jr. syn. of ornatifimbriella],; McDunnough 
1939:71 [jr. syn. of ornatifimbriella]; Busck 1939:575 [jr. syn. of ornatifimbriella]. 

Gelechia xanthuris; McDunnough 1939:72 [misident. ]. 

Filatima xanthuris; Busck 1939:576 [misident. ]. 

Gelechia amorphaeella; Chambers 1877:124; Chambers 1878a:111; Chambers 1878b:141; 
Smith 1891:100; Dyar 1903:516; Barnes & McDunnough 1917:158; McDunnough 
1939:71. 

Filatima amorphaeella; Busck 1939:575; Hodges 1983:23, [jr. syn. of ornatifimbriella]. 

Gelechia amorphelia; Busck 1903:891 [missp., unrecognized] 

Gelechia amorphella; Meyrick 1925:84 [emend.]. 


Diagnosis. Gnathos hooklike apically, costal lobe of valva produced slightly beyond 
saccular lobe, saccular lobe narrow, posterior margin of vinculum entire, lateral sclerite of 
aedeagus spinose and medially twisted, dorsal lobe truncate apically, ventral part of 
antrum with two subequal ribbonlike sclerites, posterior part of corpus bursae with dense 
microtrichia, signum small, dentate. Most specimens of ornatifumbriella can be recognized 
by the large circular discal spots and the pale grayish-brown distal 1/5 of the forewing. 

Description. Head: haustellum dark brown basally, pale grayish brown distally; maxil- 
lary palpus dark brown; dorsal and medial surfaces of labial palpus pale grayish brown, 
ventrolateral surfaces dark brown or scales pale brown basally, dark brown apically; vertex 
of second segment with individual scales pale grayish brown basally, brown or dark brown 
distally; frons with pale grayish-brown scales medially, dark-brown scales in front of eye; 
vertex and frons slightly darker than frons, scales with lustrous reflections, each scale 
gradually widened from base, apical margin entire, rounded, scales posterad of eye dark 
brown; antennal scape, pedicel, and a variable number of basal flagellomeres dark brown 
above, other flagellomeres with individual scales pale grayish brown basally, dark brown 
distally, antenna yellowish gray underneath. Thorax: mesonotum and tegula pale grayish 
brown intermixed with brown; lateral surface of legs mostly dark brown intermixed with 
pale grayish brown, tibiae with a narrow white band at 1/2 length and apex, apex of each 
tarsomere white, mesial surface white. Forewing (Fig. 1): length 6.5-8.5 mm (n = 13); 
most wing scales pale grayish brown basally, dark brown distally, each scale gradually 
widened from base, distal margin serrate (Fig. 5 of F. xanthuris); anterior margin of wing 
dark brown basally, becoming mottled dark brown and gray to 3/4 length, then pale gray- 
ish brown; a dark-brown subcircular spot at 3/5 length of cell and one at end of cell; discal 
spots large or small, equal or distal spot larger, separate or united; mid-discal spot absent 
in some specimens; scales on undersurface pale grayish brown basally, brown apically; an- 
terior margin brown. Hindwing: male with one acanthus, female with two acanthi; upper 
surface pale grayish brown, darkening slightly to apex, undersurface pale gray/off-white, 
veins and margin of wing darker; undersurface of male with curtain scaling (Figs. 8-9 of 
F. xanthuris) from wing base to slightly beyond end of cell and between Sc+R, and Rs; 
curtain scaling perpendicular to anterior margin, extending to near middle of cell; each 


VOLUME 51, NUMBER Il 35 


Fics. 1-4. Species of Filatima. 1, female F. ornatifimbriella (Clemens); 2, male F. 
xanthuris (Meyrick); 3, holotype female of F. adamsi, n. sp.; 4, holotype male of F. oc- 
cidua, n. sp. 


scale elongate, with deeply dissected distal margin; male with pale-gray sex scales be- 
tween veins posterad of curtain scaling to anal area (Figs. 8-10 of F. xanthuris). Abdomen: 
terga 1-6/7 with basal part grayish brown, distal margin white; sterna mostly pale grayish 
brown medially, brown laterally; male eighth tergum an invaginated pouch (Fig. 11), with 
piliform sex scales originating from anterolateral arms; male eighth sternum (Fig. 12) with 
two short anterolateral arms, distal emargination broad and rounded. Male genitalia (Figs. 
16, 19): gnathos elongate, narrowed distally forming a recurved hook; costal lobe of valva 
narrow throughout length, setose, extending slightly beyond saccular lobe; saccular lobes 
nearly symmetrical, setose, strongly arched medially and curved dorsally at apex; poste- 
rior margin of vinculum entire; aedeagus with lateral sclerite from zone spinose, medially 
twisted, dorsal lobe (=sclerotized lobe from zone (Hodges 1986)) truncate apically, inter- 
nal lobe longer than ventral lobe, and with broader distal margin. Female genitalia (Fig. 
22): ovipositor with two telescopic membranous parts; posterior apophyses much longer 
than anterior apophyses; membrane adjacent to anterior apophyses forming a deeply in- 
vaginated pocket bearing microtrichia; antrum with two subequal, ribbonlike sclerites 
ventrally, one subtriangular and one oval sclerite dorsally; corpus bursae and accessory 
bursae with many hairlike microtrichia within posterior end; inception of ductus semi- 
nalis on anterior part of accessory bursae; signum heavily sclerotized, small, anterior mar- 
gin finely dentate. 

Types. Gelechia ornatifimbriella Clemens. Holotype: 4; type no. 7347; “Ill[inois];” “3 
genitalia slide by RW Hodges, 2936” [Academy of Natural Sciences of Philadelphia, 
ANSP]. Gelechia unctulella Zeller. Holotype: 4; “type 1703” [red label]; “Dallas, Tex[as], 
Boll” “Zeller.” “Gelechia unctulella Z[eller]” [|handwritten, green label]; “3 genitalia slide 


36 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 5-10. Ultrastructure of scales of Filatima fore- and hindwings. 5-6, scales of 
forewing of Filatima xanthuris (Clemens); 7, scales of forewing of Filatima depuratella 
(Busck); 8-9, hindwing of male Filatima xanthuris (Clemens) showing curtain scaling 
within area from wing base to slightly beyond end of cell; 10, highly dissected male sex 
scales within lower half of discal cell, basal part of cells M3-CuA, and CuA,-CuAg, and en- 
tire anal area. Line scale = 1.0 um for Figs. 5-7, 10; line scale = 1.0 mm for Figs. 8—9. 


3285, RW Hodges.” [Museum of Comparative Zoology, MCZ]. Gelechia amorphaeella 
Chambers. Lectotype: 3; present designation; “Type 1480” [red label]; “Chambers, 
Color[ado]” “Gelechia amorphaeella, Cham|bers] Col[lection]” [handwritten label]; “Lec- 
totype, RW Hodges” [handwritten label]; “3 genitalia slide 3289, by RW Hodges.” Lecto- 
type and two paralectotypes, all with same label data, in MCZ. One specimen of syntype 
series apparently lost. 

Foodplants. Amorpha fruticosa L. (Fabaceae) (Chambers 1878a and pinned speci- 
mens). 

Distribution. Filatima ornatifimbriella is known from five localities: [Rock Island? ], 
Illinois; Halsey, Nebraska; Edgerton, Colorado; Dallas, Texas; and Riverside, California. 


VOLUME 51, NUMBER 1 27 


Fics. 11-14. Eighth tergum and sternum of Filatima. 11, eighth tergum of F. ornati- 
fimbriella (Clemens); 12, eighth sternum of F. ornatifimbriella (Clemens); 13, eighth ter- 
gum of F. xanthuris (Meyrick); 14, eighth sternum of F. xanthuris (Meyrick). Line scale 
= ilQinnen 


Adults have been reared from larvae ( June-July). The species overwinters as an adult, 
based on specimen label data; but it does not appear to have been collected at light. Spec- 
imens examined: 5 4, 8 °, § slides. 


Filatima xanthuris (Meyrick 1927) 
(Figs. YAHAS. SOM alls 17 VOL WR) 


Gelechia xanthuris; Meyrick 1927:346; McDunnough 1939:72. 

Filatima xanthuris; Hodges 1983:23 [revised status]. 

Gelechia ornatifimbriella; Clarke 1932:67, pl. 2, fig. 4 (male gen.), pl. 3, fig. 4 (female gen.) 
[misident. ]. 

Filatima ornatifimbriella; Busck 1939:575, pl. 60, Figs. 11, lla, 11b (male gen.), pl. 66, 
Fig. 44 (female gen.) [misident.]; Clarke 1969, 7:99, pl. 49, figs. 4, 4a, 4b (wing pattern, 
aedeagus, genital capsule), lectotype designation of Gelechia xanthuris Meyrick and 
treatment as junior synonym of F. ornatifimbriella [misident.]. 


38 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 15. Lateral view of male genitalia of Filatima xanthuris (Meyrick) with vinculum 
not separated from dorsal elements of genital capsule. Genital capsule rotated laterally 45 
degrees. Line scale = 1.0 mm. 


Diagnosis. Gnathos apically rounded and blunt, costal lobe of valva extending well be- 
yond saccular lobe, inner surface of left saccular lobe with a broad and angular basal lobe, 
right saccular lobe slightly widened basally, posterior margin of vinculum slightly emar- 
ginate medially, aedeagus with lateral sclerite from zone fingerlike, dorsal lobe obtuse api- 
cally, internal lobe slightly longer and narrower than ventral lobe. Most specimens of F. 
xanthuris can be recognized by the united, subrectangular discal spots of the forewing. 

Description. As for F. ornatifimbriella except: Forewing (Fig. 2): length 6.0—10.1 mm 
(n = 208); individual scales pale brownish gray basally, brown or dark brown distally; dis- 
cal spots variable, usually subrectangular, large or small, subequal or unequal, separate or 
united; many specimens with various spots and/or streaks basal to spot at middle of cell; 
two short streaks along CuP (united in some specimens), one basal spot on midline of cell; 
one subcostal spot between spots at base of middle of cell; basal spots appear to be pres- 
ent or absent in any combination. Abdomen: male eighth tergum narrow (Fig. 13); distal 
margin of male eighth sternum deeply emarginate medially (Fig. 14). Male genitalia (Figs. 
15, 17, 20): gnathos apically rounded and blunt; costal lobe of valva narrow throughout 


VOLUME 51, NUMBER 1 39 


Fics. 16-18. Male genitalia of Filatima. 16, male genitalia of F. ornatifimbriella 
(Clemens); 17, male genitalia of F. xanthuris (Meyrick); 18, male genitalia of F. adamsi, 
n. sp. Line scale = 1.0 mm. 


40 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


length, setose, extending well beyond saccular lobe; saccular lobes asymmetrical, inner 
surface of left lobe with a broad and angular lobe, right lobe widened basally; posterior 
margin of vinculum slightly emarginate medially; aedeagus with lateral sclerite from zone 
fingerlike; apical margin of dorsal lobe obtuse; internal lobe slightly longer and narrower 
than ventral lobe. Female genitalia (Fig. 23): as for F. ornatifimbriella, except lobe adja- 
cent to anterior apophyses slightly invaginated; antrum membranous. 

Types. Lectotype: 4, designated and figured by Clarke (1969), BM slide no. 5771, Div- 
idend, Utah, 26 April. Lectotype and nine paralectotypes in The Natural History Mu- 
seum, London, BM(NH). 

Foodplants. Thermopsis pinetorum Greene, Lupinus sp., Robinia spi}, Viciad_ sp: 
(Fabaceae) (pinned specimens). 

Distribution. Filatima xanthuris has been collected in the mountains of North and 
South Carolina; the Boston Mountains, Arkansas, Tenkiller Lake, Oklahoma; Silverton, 
Colorado; Guadalupe Mountains, Texas; Lincoln County, New Mexico; Coconino County, 
Arizona; Utah County, Utah; Lander County, Nevada; Riding Mountains, Manitoba; 
Rocky Mountains in Alberta and British Columbia; generally in Washington; and Lincoln 
County, Oregon. Adults have been collected from 26 March to 10 October. Specimens ex- 
amined: 96 3, 108 °, 68 slides. 

Remarks. Preliminary, comparative studies of the ultrastructure of the dorsal 
forewing scales of F. xanthuris and F. depuratella (Busck) indicate that Filatima with 
shiny scales have large windows between the longitudinal ridges of each scale, whereas 
Filatima with dull scales have either few smaller windows or no windows between longi- 
tudinal ridges (Figs 6-7). 


Filatima adamsi Hodges & Adamski, new species 
(Figs. 3, 18, 21) 


Diagnosis. Frons with scales in front of eye generally concolorous with rest of frons, a 
few dark-brown scales present. Gnathos broadly curved, costal lobe of each valva slender, 
gradually widening from 2/5 length to apex, extending well beyond saccular lobe, saccular 
lobes of valvae asymmetrical, lobe of left valva with mesial margin broadly curved, lobe of 
right valva with mesial margin sinuous, becoming narrower apically; posterior margin of 
vinculum entire; aedeagus with sclerites as illustrated. Female genitalia unknown. 

Description. Head: haustellum and maxillary palpus grayish brown basally, becoming 
pale grayish brown distally; labial palpus with first segment mainly dark gray brown later- 
ally, second segment grayish orange dorsobasally, then brown almost to apex, apex and 
ventral scale tuft mottled pale and dark gray, scale tuft narrow, third segment mainly dark 
brown with scattered off-white scales; frons, vertex, and occiput mainly shining yellowish 
gray, individual scales darker at apices, scales on frons tipped with dark brown, several 
brown scales in front of eye, scales on vertex and occiput with small gray area at apex of 
each; scape and shaft of antenna mainly dark brown, ventral surface of scape yellowish 
gray, individual scales on shaft paler at base than apex, antenna broken after flagellomere 
10/11; ocellus present. Thorax: mesonotum and tegula appearing gray brown, individual 
scales pale yellowish gray basally, gray brown distally; foreleg coxa mottled yellowish gray 
and gray brown, mainly dark, femur, tibia, and tarsus medium to dark gray brown, with 
pale scales just beyond 1/2 length of tibia and at apex, apex of tarsomeres 1—4 off-white; 
midleg similar to foreleg but slightly darker; hindleg unknown. Forewing (Fig. 3): length 
7.0 mm (n = 1); mainly dark gray brown, individual scales paler at base; an irregular dark 
gray-brown mark at 3/5 length of cell, a dark gray-brown blotch at end of cell, several dark 
gray-brown scales along fold from near base to 3/4 length of fold, several pale-gray scales 
on anterior and posterior margins at 4/5 wing length, fringe on distal margin mainly pale 
gray, individual scales tipped darker gray. Hindwing: undersurface with curtain scales, ad- 
ditional sex scales extending from base of wing toward posterior margin and in the cell to 
vein 2A. Abdomen: upper surface with mostly pale grayish-brown scales intermixed with 
brown scales; undersurface mostly white mesially, mostly dark-brown scales intermixed 
with brown scales ventrolaterally; male eighth tergum most similar to that of xanthuris, 
sternum most similar to that of ornatifimbriella, except posterior margin less emarginate. 


VOLUME 51, NUMBER 1 4] 


Fics. 19-21. Male genitalia (aedeagi) of Filatima (for orientation note that the ductus 
ejaculatorius enters the aedeagus on the dorsal surface). 19, aedeagus of F. ornatifim- 
briella (Clemens); 20, aedeagus of F. xanthuris (Meyrick); 21, aedeagus of F. adamsi n. 
sp. Line scale = 1.0 mm. 


42 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Male genitalia (Figs. 18, 21): vinculum asymmetric, saccus directed toward right; costal 
lobe of valva wider distally than at base, extending well beyond saccular lobe; saccular 
lobes asymmetrical, left lobe broad, right lobe narrower apically; posterior margin of vin- 
culum entire; gnathos hook shaped, strongly curved in distal 1/3; aedeagus with several 
sclerotized flanges and plates on distal 1/2; a prominent, subtriangular plate with serrate 
lateral margin. Female genitalia: unknown. 

Types. Holotype: 3. Label data: “M[aine]: West P[oint], Little Wood Is[land], 13 
August] 1972, S. B. Adams leg.” “d Genitalia slide by DA, USNM 87529” [green label]. 
Holotype in National Museum of Natural History, USNM. 

Foodplants. Unknown. 

Distribution. Filatima adamsi is known only from the type locality. It was collected at 
incandescent light. 

Remarks. This species appears closely allied to F. vaniae Clarke by sharing similarly 
shaped costal lobes of the valvae and the aedeagus with a subtriangular ventral sclerite 
with a serrate edge. They differ in that adamsi has wider saccular lobes and longer costal 
lobes of the valvae. In addition, vaniae is paler. The hindlegs are missing on the holotype. 

Etymology. This species is named after its collector, Mrs. Sally B. Adams (now Mrs. 
S. A. Brady). 


Filatima occidua Hodges & Adamski, new species 
(Figs. 4, 24) 


Diagnosis. A small medium to dark-gray moth with a small dark-brown spot at 2/3 the 
length of the cell and an irregular dark-brown mark at the end of the cell on the forewing. 
Dorsodistal margin of antrum heavily sclerotized and broadly incurved mesially; antero- 
lateral margin of seventh abdominal sternum with a V-shaped invagination. Right lobe of 
corpus bursae with dense, fine spicules on basal 1/3; zone of spicules extending basome- 
sially onto antrum. Signum with a pair of inwardly directed, triangular, lateral lobes; each 
lobe dentate, particularly on posterior margin. 

Description. Head: maxillary palpus and base of haustellum dark brown, haustellum 
becoming yellowish brown by 1/2 length; labial palpus mainly brown, individual scales 
pale gray brown at extreme base of each scale, dorsomesial surface of second segment 
pale gray from base to 1/2—3/5 length, third segment mottled with many pale-gray/off- 
white scales, particularly on posterior margin; antenna mottled dark brown and pale 
gray/off-white, ventral surface of scape pale, most scales of shaft pale nearly to apices, dis- 
tal margins very dark brown; frons, vertex, and occiput mainly shining pale yellowish gray, 
dark-brown scales in front of eye directed ventromesially and slightly overlapping on ven- 
tral part of frons, scales on vertex and occiput very narrowly margined with slightly darker 
gray; ocellus present. Thorax: tegula dark brown at base, yellowish gray distally, individual 
scales narrowly margined with darker gray, mesothorax mainly medium to pale gray, indi- 
vidual scales tipped and streaked darker gray; foreleg with coxa mottled dark brown and 
pale gray, individual scales streaked darker gray on distal 3/4, femur, tibia, and tarsus 
mainly brown, individual scales pale gray based, apex of each tarsomere with off-white 
scales; midleg similar to foreleg but surfaces generally paler, tarsomeres with many gray 
scales; hindleg as for midleg, mesial surface noticeably very pale gray/off-white. Forewing 
(Fig. 4): length 5.5-6.2 mm (n = 5): mainly dark gray, yellowish-brown tipped scales on 
basal 1/5, dark-gray tipped scales from 1/5—4/5 length and dark-brown tipped scales at 
apex, a brown spot at 3/5 length of cell and dark-brown blotch at end of cell. Male geni- 
talia: unknown. Female genitalia (Fig. 24): salient features given in diagnosis. : 

Types. Holotype: 2, “Pullman, W[ashingto]n, J.F. Clarke, 3-VITI-[19]32”; “Reared from 
Lupinus ornatus [Douglas x Lindl.]”; “3144”; “2 genitalia slide by AB, USNM 9716” [green 
label]; “2 genitalia slide by AB, Aug[ust] 30/[19]43” [handwritten label]. USNM. 
Paratypes: 3 °, “Mill Valley, Marin Co[unty], Callifornia], IX-26-1925”; “H.H. Keifer, Col- 
lector”; “2 genitalia slide by DA, USNM 87551” [green label]; “° genitalia slide by DA, 
USNM 87519” [green label]; one specimen not dissected; °, same data as above except, 
“February 4, 1926” °, “July 24-31”. Paratypes in California Academy of Sciences, San 
Francisco and USNM. 


VOLUME 51, NUMBER Il 43 


Fics. 22-23. Female genitalia of Filatima. 22, female genitalia of F. ornatifimbriella 
(Clemens); 23, female genitalia of F. xanthuris (Meyrick). Line scale = 1.0 mm. 


4 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 24. Female genitalia of Filatima occidua, n. sp. Line scale = 1.0 mm. 


VOLUME 51, NUMBER 1 45 


Foodplants. Lupinus sericeus Pursh, var. sericeus (Fabaceae). Currently, L. ornatus 
Douglas x Lindl. is a junior synonym of L. sericeus Pursh, var. sericeus. 

Distribution. Fatima occidua is known from southeastern Washington and Marin 
County, California. Adults have been collected on 4 February, 3 August, and 26 Septem- 
ber. 

Etymology. The specific epithet is derived from the Latin occiduus, -a, -um, meaning 
setting [of the sun] and referring to the distribution of occidua. 


DISCUSSION 


Comparison of the male and female genitalia suggest that the Fila- 
tima treated herein probably are distantly related and do not represent 
a monophyletic group within the genus. The lustrous appearance of the 
vestiture shared by these species probably is a homoplastic feature 
within the genus. The shape of the aedeagus of F. xanthuris is highly un- 
usual within Filatima and immediately separates it from the other spe- 
cies. Until a phylogenetic analysis is completed for all Filatima, a suit- 
able hypothesis of relationships among species cannot be made. 


ACKNOWLEDGMENTS 


We thank Susann G. Braden and Walter R. Brown of the Scanning Electron Mi- 
croscopy Laboratory, Smithsonian Institution, for help with our ultrastructural studies; 
Laurence J. Dorr, Department of Botany, Smithsonian Institution, for providing taxo- 
nomic information on hostplants; and Philip D. Perkins, Museum of Comparative Zool- 
ogy, Harvard University, for the loan of the holotype of Gelechia unctulella Zeller, and 
syntypes of G. amorphaeella Chambers. We also thank Richard L. Brown, Donald R. 
Davis, Douglas C. Ferguson, Natalia J. Vandenberg, and two anonymous reviewers for 
their helpful comments. 


LITERATURE CITED 


BARNES, W. & J. MCDUNNOUGH. 1917. Check list of the Lepidoptera of boreal America. 
Herald Press, Decatur, Illinois. ix+392 pp. 

Busck, A. 1903. A revision of the American moths of the family Gelechiidae. Proc. U. S. 
Natl. Mus. 25(1304):767—926. 

. 1939. Restriction of the genus Gelechia (Lepidoptera: Gelechiidae), with descrip- 
tions of new genera. Proc. U.S. Natl. Mus. 86(3064):563—593, pls. 58—71. 

CHAMBERS, V. T. 1877. The Tineina of Colorado. Bull. U. S. Geol. Geogr. Surv. Territ. 
3(1):121-142. 

. 1878a. Tineina and their food-plants. Bull. U. S. Geol. Geogr. Surv. Territ. 

4:107-123. 

. 1878b. Index to the described Tineina of the United States and Canada. Bull. U. 
S. Geol. Geogr. Surv. Territ. 4:125—167. 

CLARKE, J. F. G. 1932. New microlepidoptera from the Pacific coast; (Gelechiidae). Can. 
Entomol: 64:63-—69. 

. 1941. The preparation of slides of the genitalia of Lepidoptera. Bull. Brooklyn 

Entomol. Soc. 36:149-161. 

. 1947. Notes on, and new species of, American moths of the genus Filatima Busck 

(Gelechiidae: Lepidoptera). J. Washington Acad. Sci. 37(8):263—275. 

. 1969. Catalogue of the type-specimens of microlepidoptera in the British Mu- 
seum (Natural History) described by E. Meyrick. Vol. 7. Trustees British Museum 
(Nat. Hist.), London. 531 pp. 

CLEMENS, B. 1864. North American microlepidoptera. Proc. Entomol. Soc. Philadelphia 
2:415—430. 


46 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Dyar, H. G. [1903] 1902. A list of North American Lepidoptera and key to the literature 
of this order of insects. U. S. Natl. Mus. Bull. 52:xix+723 pp. 

FORBES, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Cornell 
Univ. Agric. Exp. Sta. Mem. 68:1—729. 

HODGES, R. W. 1983. Gelechiidae. In R. W. Hodges et al. (eds.), Check list of the Lepi- 
doptera of America north of Mexico. E. W. Classey Ltd. and The Wedge Entomol. 
Res. Found., London. 

. 1986. Gelechioidea: Gelechiidae (in part). In Dominick, R.B., et al. (eds.), The 
moths of America north of Mexico, Fasc. 7.1. 

HUEMER, P. 1987. Eine modifizierte Genitalpraparationstechnik fiir die Gattung Cary- 
ocolum. Mitteil. Schweiz. Entomol. Ges. 60:207—211. 

Kiots, A. B. 1970. Lepidoptera. In Tuxen, S. L. (ed.), Taxonomist’s glossary of genitalia 
in insects. 2nd ed. Munksgaard, Copenhagen. 

KORNERUP, A. & J. H. WANSCHER. 1978. Methuen handbook of colour. 2nd ed. Methuen 
and Ce., Ltd., London. 243 pp. 

McDUNNOUGH, J. 1939. Checklist of the Lepidoptera of Canada and the United States of 
America. Part II. Microlepidoptera. Mem. So. California Acad. Sci. 2(1):1-171. 

MEYRICK, E. 1925. Lepidoptera Heterocera fam. Gelechiadae. In Wytsman, P. (ed.), Gen- 
era insectorum, 184:1—290, pl. 1-5. 

. 1927. Exotic Microlepidoptera 3(11):321—352. 

PITKIN, L. 1984. A technique for preparing complex male genitalia in microlepidoptera. 
Entomol. Gaz. 37:173-179. 

SATTLER, K. 1960. Generische Gruppierung der europaéischen Arten der Sammelgattun 
Gelechia (Lepidoptera, Gelechiidae). Deutsche Entomol. Zeitschr. N. F. 7(1/2): 
10-118. 

SMITH, J. B. 1891. List of the Lepidoptera of boreal America. Am. Entomol. Soc., 
Philadelphia. vi+ 124 pp. 

ZELLER, P. C. 1873. Beitraége zur Kenntniss der nordamericanischen Nachtfalter, beson- 
ders der Microlepidopteren. Zweite Abtheilung. Verhandl. Zool.-Bot. Ges. Wein 
23:201—334, pls. III-IV. 


Received for publication 12 August 1995; revised and accepted 2 F. ebruary 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 47-56 


AN EXAMPLE OF CLINAL VARIATION IN EASTERN NORTH 
AMERICAN BUCKMOTHS (SATURNIIDAE: HEMILEUCA) 


BRIAN G. SCHOLTENS 
Biology Department, College of Charleston, Charleston, South Carolina 29424, USA 


AND 


WARREN HERB WAGNER, JR. 
Department of Biology, University of Michigan, Ann Arbor, Michigan 48109, USA 


ABSTRACT. Morphological variation in populations of eastern North American 
buckmoths was examined. Samples of 25 males each were analyzed from four points along 
a north-south line approximately 800 km long from Schoolcraft Co., Michigan to Vinton 
Co., Ohio. Clinal variation was demonstrated from higher to lower latitudes: (1) forewing 
length from smaller to larger; (2) forewing white band width from wider to narrower; and 
(3) darkness of the black background of the wing from lighter to darker. No logical way 
was found to separate these four populations into subspecies or species based on morpho- 
logical characters. 


Additional key words: Hemileuca maia, H. nevadensis, H. lucina. 


The black and white buckmoths of the eastern United States and 
Canada have attracted much attention because of their showy wing pat- 
terns and unusual flight period very late in the season. Three species 
are usually attributed to eastern North America: in the western part 
Hemileuca nevadensis Stretch, in the northeast H. lucina Hy. Edw., and 
in the remainder of the area H. maia (Drury) (Ferguson 1971, Covell 
1984). Each species is generally associated with a distinct food plant, all 
unrelated: willows (Salix: Salicaceae), meadowsweets (Spiraea: Rosa- 
ceae), and oaks (Quercus: Fagaceae), respectively. 

We have surveyed various habitats and have verified, by egg mass 
placement and larval feeding, a more diverse range of food plants than 
is usually assumed (Scholtens & Wagner 1994). In addition, several pop- 
ulations are difficult to assign to any of the three recognized North 
American species. Some appear to be intermediate in maculation be- 
tween H. nevadensis and H. lucina and others between H. nevadensis 
and H. maia. Here, using data on the characters typically used to distin- 
guish the presently recognized species, we test the hypothesis that the 
populations in the Great Lakes region consist of a single species form- 
ing a cline ranging from relatively small, ‘washed-out’ looking forms in 
the north to larger, heavily marked forms in the south. 


MATERIALS AND METHODS 


We sampled and studied buckmoths at four localities in the Great 
Lakes region. From north to south these were: (1) Schoolcraft County 
in the Upper Peninsula of Michigan; (2) Roscommon County in the 


48 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


™ 
Seserertt | corey = %, 
—— 
ice 
} 


> 
AS SINE 


i ee 


i 
Dame) 7 7 
fp ALPS 


fe ere 


Fic. 1. Map of sampling localities and example of male buckmoth from each. 


central Lower Peninsula; (3) Washtenaw County in the southern Lower 
Peninsula; and (4) Vinton County in southern Ohio (Fig. 1). At each lo- 
cality we made notes on the habitat and host plants used by the buck- 
moths and collected a series of 40—50 males, from which a sample of 25 
in good condition were chosen for measurements and visual assessment. 
Vouchers from each population have been deposited at the University 
of Michigan Museum of Zoology. 

A video image of each specimen was stored on a Macintosh II com- 
puter using the program NEH Image version 1.26. The lighting ar- 
rangement and the specimen-camera distance were not changed during 
image capture. For each image, measurements were taken, using Image, 
of forewing length from the base of the wing to the farthest point on the 
wing tip, and width of the white band on the forewing and hindwing 
along veins M, and CU,. The limits of the white band were easily deter- 
mined by a sharp change over in scale color, even in the most diffusely 


VOLUME 51, NUMBER 1 49 


patterned moths. In addition, 10 pixel by 10 pixel areas were marked off 
on the forewing between veins CU, and CU, inside the white band and 
immediately distal to the white band. Using Image, an average darkness 
for each 100 pixel area was calculated based on a 256 point gray scale. 

Because one characteristic difference between the populations was 
wing darkness, we did a microscopic examination of scale sizes and 
densities for the three Michigan populations, covering most of the vari- 
ation in darkness. For 10 specimens from each population, we counted 
all scales in a 1.42 x 2.16 mm area of the forewing between veins M, 
and CU,, centered on the white band. In addition for 5 specimens 
from each population, 10 scales in one field of view were measured for 
length and width. For all data, comparisons among localities were 
made by ANOVA using the statistical package SYSTAT 5.0 for the 
Macintosh. 


RESULTS 


A great deal of variation exists in the habitats and host plants of Great 
Lakes buckmoths compared to previously published data. In the north- 
ern part of the range through southern Michigan, habitats are mainly 
wetlands, similar to those typically used by the western H. nevadensis, 
while in the south, upland wooded areas are prevalent. The host plants 
in wetland areas include willow, poplar, meadowsweet, and bog birch, 
and in the dry wooded sites, oaks (Scholtens & Wagner 1994). 

Forewing length in the sampled populations varied from a mean of 
24.04 mm in northern Schoolcraft County to 26.59 mm in southern Vin- 
ton County (Fig. 2). ANOVA shows that forewing length increases sig- 
nificantly from north to south (F = 20.30, P = <0.001). The width of the 
white band on the forewing decreases significantly from north to south 
(on M,: F = 40.96, P = <0.001; on CU,: F = 24.47, P = <0.001). On vein 
M, the mean width is 2.17 mm in Schoolcraft County and 0.61 mm in 
Vinton County. On vein CU, the mean width is 4.42 mm in Schoolcraft 
County and 2.60 mm in Vinton County (Fig. 3). Although the width 
of the white band on the hindwing varies significantly among the 
three populations (on M;: F = 2.79, P = 0.045; on CU,: F = 15.30, P = 
<0.001), we saw no demonstrable trend from north to south (Fig. 4). 

There is a significant increase in the darkness of the white band from 
north to south (F = 5.04, P = 0.003), varying from Schoolcraft County 
with a mean of 82.97 gray scale units (out of 256) to Vinton County at 
88.19 units. There is a much more pronounced significant increase in 
the darkness of the black areas of the wing from north to south (F = 
142.28, P < 0.001), varying from 112.34 units in Schoolcraft County to 
164.12 units in Vinton County (Fig. 5), with most of the change occur- 
ring from northern to southern Michigan. These means hide a great 


50 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Forewing Length (mm) 


Schoolcraft Roscommon Washtenaw Vinton 


Location (North to South) 


Fic. 2. Mean forewing length of sampled populations (error bars indicate standard 
deviation). 


deal of variation in the darkness of the black areas of the wing, enough 

so that wing darkness overlaps in all adjacent populations (Fig. 6). 
Darker wing color in the southern buckmoth populations is due to 

the size and distribution of wing scales. There are substantial differ- 


Ee 
2 
= 
TG 
> Mi FORECU2 
3 fH] FOREMi1 
ci 
a 


Schoolcraft Roscommon Washtenaw Vinton 


Location (North to South) 


Fic. 3. Mean width of white band on forewing veins CU, and M, (error bars indicate 
standard deviation). 


VCLUME 51, NUMBER 1 51 


M@ HIND Cu2 
Ea HIND M1 


Band Width (mm) 


Schoolcraft Roscommon Washtenaw Vinton 
Location (North to South) 


Fic. 4. Mean width of white band on hindwing veins CU, and M, (error bars indicate 
standard deviation). 


ences in both size and density of scales among the populations (Figs. 7 
and 8). From north to south, scale length (F = 23.97, P = <0.001), width 
(F = 7.30, P = 0.008) and density (F = 26.97, P = <0.001) increase sig- 
nificantly. All three of these parameters contribute to the lighter, more 
translucent appearance of the wings in the north. 

Increasing forewing band width with decreasing wing length accen- 
tuates the trend of lighter colored wings in the north, and in Vinton 
County the ratio of band width to wing length is half or less of that in 


200 


M™ DARK AREA 
& LIGHT AREA 


100 


Gray Scale Value 


Schoolcraft Roscommon Washtenaw Vinton 
Location (North to South) 


Fic. 5. Mean darkness of light and dark areas of forewing (error bars indicate stan- 
dard deviation). 


52 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Schoolcraft Co. 


Number of Specimens 


Roscommon Co. 


Number of Specimens 


Washtenaw Co. 


Number of Specimens 


Number of Specimens 


(2) 
© 
= 


Se 
an 7 8 
Paes NS 

o 


115-120 
125-130 
135-140 
165-170 
170-175 
175-180 
180-185 
185-190 


145-150 
160-165 


Ww 
“2) 
- 
1 
jo} 
w 
= 


100-105 
105-110 
110-115 
120-125 
130-135 
140-145 


iB 
Density (on 256 pt. gray scale) 


Fic. 6. Distribution of wing darkness at the four study sites. 


Schoolcraft County. Also influencing this impression is a proportionally 
greater increase in width at the center of the white band as you proceed 
north. The ratio of the width of the white band on vein M, to the width 
on vein CU, decreases to less than half of the Schoolcraft County value 
in Vinton County. 


VOLUME 51, NUMBER Il 53 


(micrometer units) 


Mean 


14 


—_ 
N 


=x 
Oo 


Lengih 
Fa Width 


Schoolcraft Roscommon Washtenaw 


Location (North to South) 


Fic. 7. Mean scale length and width (error bars indicate standard slevtetion). 


Mean (scales/square mm) 


Bre. 8: 


120 


100 


80 


60 


40 - 


20 


Schoolcraft Roscommon Washtenaw 
Location (North to South) 


Mean scale density (error bars indicate standard deviation). 


o4 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


DISCUSSION 


Our results clearly demonstrate correlated changes in food plants and 
maculation characters of buckmoths along a north-south line in the 
Great Lakes region. Where the wing length is smaller in the north, the 
white band is wider and the black background is paler. The northern- 
most element is most similar to the New England buckmoth, Hemileuca 
lucina, the southernmost element is most similar to H. maia, and the in- 
termediates resemble some of the forms of the widespread western 
buckmoth, H. nevadensis. There is also a general correlation with habi- 
tat, bogs and fens in the north and upland, oak woods in the south. The 
larval food plants include bog birch (Betula pumila L.: Betulaceae), wil- 
lows (Salix spp.) and meadowsweets (Spiraea spp.) northward and oaks 
(Quercus spp.) in the south (Scholtens & Wagner 1994). 

Change in maculation of buckmoths is a direct result of a change in 
scale size and density. Scale density in the northern populations de- 
creases, even when corrected for the reduction in wing size, and scale 
size also decreases proportionally much more than wing size. The exis- 
tence of a north-south morphological cline in Hemileuca should not be 
surprising. Good examples of latitudinal clines exist in other widespread 
species in North America (e.g., Cercyonis, Emmel 1969), Great Britain 
(e.g., Coenonympha, Porter 1980), and Australia (e.g., Tisiphone, Lucas 
1969). In our experience, forms of non-migratory species in the extreme 
north tend, in general, to be smaller and the pattern less contrasting 
than those from the south. The same is often true of spring versus sum- 
mer forms. 

Some speculations about possible evolutionary causes of size and mac- 
ulation changes in northern populations are the following: (1) smaller 
size may simply be a result of a shorter growing season; and (2) scale de- 
velopment may be aborted because of the shorter pupal period in 
northern populations (late June through late August in the north versus 
mid-June through early October in the south). These remain specula- 
tions until experimental work can verify or refute them. 

Confusion about the nomenclature of Hemileuca populations has ex- 
isted for many years as evidenced by the naming of H. latifascia Barnes 
and McDunnough as a subspecies of H. lucina, followed by its synonomi- 
zation with H. nevadensis (Ferguson 1971). Forbes (1960) even remarked 
on the intermediacy of H. latifascia between H. maia and H. lucina. Great 
Lakes populations have always been vexing because willow feeders have 
periodically been reported from the region (Ely 1954, Riley 1873, Wor- 
thington 1878), but not studied carefully. Our data show that no good 
maculation differences exist that allow Great Lakes populations to be 
placed confidently into one of the recognized species. Likewise, the host 


VOLUME 51, NUMBER Il 55 


plants and habitats are not distinctive and do not serve to identify pop- 
ulations definitively as once thought. Ferge (1981) documented similar 
within population variation in Wisconsin, and populations in Minnesota, 
and the eastern United States also resemble those we studied. Legge et 
al. (1996), in a study examining allozyme differences in Hemileuca popu- 
lations from across the country, found very few differences between any 
of the populations examined, and suggested that these populations could 
still be considered distinct species based on ecological differences. Our 
data would not support this, showing that ecological differences among 
the various populations are not consistent (Scholtens & Wagner 1994). 
Although separate eastern populations may seem quite distinct, local 
population differentiation may be more prevalent in this region because 
of longer isolation due to habitat fragmentation. 

The patterns seen in this study could result from variation within a 
single species or from a hybrid zone between distinct species. Our data 
cannot distinguish between these alternatives, but the most satisfying 
explanation for our findings is that all populations represent a single 
species showing clinal variation from north to south. A species distinc- 
tion could exist between the wetland populations that feed on birch, 
poplar, meadowsweet and willow and the upland, oak-feeding popula- 
tions. Ecologically, these appear distinct and some evidence indicates 
that they may be isolated by pheromone differences (James Tuttle, 
pers. comm.), but these differences are not indicated by the clinal na- 
ture of the morphological variation. If all populations are a single spe- 
cies they would be referred to as H. maia (Drury). If two entities exist, 
the upland populations would be H. maia (Drury) and the wetland ones 
H. nevadensis Stretch. This question may be decided conclusively by 
careful hybridization and rearing studies between geographically close 


populations of both types. 


ACKNOWLEDGMENTS 


We were supported by the Michigan Non-game Wildlife Fund (Department of Natural 
Resources). The staffs of the University of Michigan Biological Station and Seney Na- 
tional Wildlife Refuge allowed us to study populations in northern Michigan. The Ameri- 
can Museum of Natural History and The University of Michigan Museum of Zoology 
loaned specimens for comparison. Specimens were also examined at Michigan State Uni- 
versity (East Lansing), the Canadian National Collection (Ottawa), the Milwaukee Public 
Museum (Milwaukee) and the Carnegie Museum (Pittsburgh). Mark O’Brien made avail- 
able the video camera for image capture and Barry O’Connor graciously allowed the use 
of his computer. David Bay, John Cryan, Robert Dirig, Les Ferge, J. Donald Lafontaine, 
Barbara Madsen, Robert Masta, Eric Metzler, Mogens Nielsen and Heidi Appel all con- 
tributed to the success of the project. 


LITERATURE CITED 


COVELL, C. V., JR. 1984. A field guide to the moths of eastern North America. Houghton 
Mifflin Co., Boston. 496 pp. 


36 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ELy, R. 1954. Concerning Hemileuca maia in Wisconsin. Lepid. News 8:29. 

EMMEL, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis (Satyri- 
dae). I. Characteristics of the genus. J. Lepid. Soc. 23:165—-195. 

FERGE, L. A. 1981. Observations on the buck moth in Wisconsin. News. Wisc. Entomol. 
Soc. 9(1): 2-5. 

FERGUSON, D. C. 1971. Bombycoidea (in part), pp. 101-153. In Dominick, R. B. et al. 
(ed.), The moths of America north of Mexico, Fascicle 20.2A. E. W. Classey, London. 

ForBES, W. T. M. 1960. Lepidoptera of New York and neighboring states. Part 4. Agaris- 
tidae through Nymphalidae including butterflies. Cornell Univ. Agric. Exp. Sta. 
Mem. 371:1—188. 

LEGGE, J. T., R. RousH, R. DESALLE, A. P. VOGLER & B. May. 1996. Genetic criteria for 
establishing evolutionarily significant units in Cryan’s buckmoth. Cons. Biol. 
10:85—98. 

Lucas, A. M. 1969. Clinal variation in pattern and colour in coastal populations of the 
butterfly Tisiphone abeona (Donovan) (Lepidoptera: Satyrinae). Aust. J. Zool. 
17:37—48. 

PORTER, K. 1980. A quantitative treatment of clinal variation in Coenonympha tullia 
(Miiller) (Lep., Satyridae). Entomol. Monthly Mag. 116:71—82. 

RILEY, C. V. 1873. Report of the noxious, beneficial, and other insects of the state of Mis- 
souri, 5. 

SCHOLTENS, B. G. & W. H. WAGNER, JR. 1994. Biology of the genus Hemileuca (Lepi- 
doptera: Saturniidae) in Michigan. Great Lakes Entomol. 27:197—207. 

WORTHINGTON, C. E. 1878. Miscellaneous memoranda. Can. Entomol. 10:15—16. 


Received for publication 23 January 1995; revised and accepted 28 January 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 57-61 


A NEW SPECIES OF RIODINIDAE FROM COLOMBIA 


CURTIS J. CALLAGHAN 
Avenida Suba 130—25, Casa 6, Bogota, Colombia 


AND 


JULIAN SALAZAR 


Museo de Historia Natural, Universidad de Caldas, AA.275, Manizales, 
Caldas, Colombia 


ABSTRACT. A new riodinid species, Calydna volcanicus, from Cerro Aguacatal 
and Cerro Clavijo in the western department of Caldas, Colombia, is described and illus- 
trated. Comments on its habitat and adult behavior are presented, with a list of other Ri- 
odinidae found in the same habitat and a range extension for Amphiselenis chama 
(Staudinger 1888). 


Additional key words: neotropical South America, coffee plantations. 


Forests at altitudes between 1300 and 2200 m in the Colombian An- 
des are fertile for the discovery of new and interesting riodinid butter- 
flies (Callaghan 1983, Salazar & Constantino 1993, Salazar, 1993). An 
unusual habitat for riodinids described in this paper consists of coffee 
plantations, considered to be an example of extreme habitat alteration. 
In these situations, the original forest has been completely destroyed 
except for some of the taller trees that have been left to shade the cof- 
fee. If insecticides have not been used extensively, this altered habitat 
can support a diverse fauna. 

The purpose of this paper is to describe a new species of Calydna, 
and present comments on the coffee zone habitat and some of the other 
riodinid species found there. The description is based on material col- 
lected by J. Escobar at the localities noted below. Measurements were 
done with a caliper, and the genitalia were prepared by soaking in 10% 
KOH solution and examined under a binocular microscope. Terminolgy 
for the genitalia follows Klots (1970) and that for veins and cells follows 
the Comstock-Needham system (Miller 1969). The citing of Riodinidae 
as a family instead of a subfamily follows Harvey (1987). 


Calydna volcanicus Callaghan & Salazar, new species 
(Figs. 1-7) 


Description. Eyes brown, not hairy, surrounded by white scaling; palpi dimorphic, 
those of female longer than male; antenna length 6.4 mm, 0.59 length of forewing; shaft 
with white scales between segments, club flat, spoon shaped; forewing costa straight, apex 
pointed, distal margin scalloped at the end of cell M,-CU,; hindwing with apex rounded, 
margin scalloped between CU2 and 2A; anal angle pointed, inner margin straight; fringe 
light brown. 

Male (Figs.1, 2): forewing length of holotype 13 mm, range of material examined 11-13 
mm (n = 3). Thorax ventrad and appendages with long, white hairs; abdomen brown with 


58 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-7. Calydna volcanicus, new species. 1, male holotype, dorsal surface; 2, same, 
ventral surface; 3, male genitalia, caudal view; 4, same, lateral view; 5, female, dorsal sur- 
face; 6, same, ventral surface; 7, female genitalia. 


VOLUME 51, NUMBER 1 59 


long, white hairs ventrad on first 5 segments; long, lateral scent hairs on last segment. 
Dorsal surface of wings dark brown with faint black and orange maculations and black 
distal margins. Forewing with two faint black parallel lines at end of cell, a single spot be- 
low the cell between CU, and 2A and faint disperse orange scaling in discal area between 
CU, and 2A. Hindwing with 2 faint parallel black lines beyond end of cell and long dis- 
perse scent hairs in discal area. Ventral surface light brown with striated dark brown mac- 
ulations. Forewing with two large black spots beyond end of cell; one in cell CU, and 
three in cell CU,. Hindwing with numerous black maculations. Genitalia (Figs. 3, 4): with 
socii unlobed; gnathos simple, unmodified; vinculum narrow, slightly bowed halfway down 
each side, saccus small, pointed; valvae broad with two protrusions, dorsad and basad, the 
dorsal protrusion longer and curving inward; transtilla broad, rounded caudad; annellus 
with a long, pointed process curving caudad; aedeagus with the tip forming a broad plate 
with teeth projecting caudad. 

Female (Figs. 5, 6): forewing length 13.0-13.5 mm (n = 2). Body dark brown dorsad, 
white ventrad. Dorsal surface of wings dark brown and yellow-orange with black macula- 
tions. Forewing with discal area between M, and 2A orange-yellow, extending to M, past 
end of cell, and with some scattered orange-yellow scaling within cell; two black spots in 
cell M, beyond cell and two in cell CU, below discal cell. Hindwing with large irregular 
yellow-orange spot in discal area between R, and cell 2A; limbal area with faint, irregular 
dark brown spots along margin between the veins. Ventral surface light brown and light 
orange, with lighter brown maculations. Forewing submargin white with brown macula- 
tions; discal area with light yellow spot as in dorsal surface; three black spots in discal cell, 
one in cell CU, and three in cell CU,. Hindwing distal half white with numerous dark 
brown striated maculations; basal half same but with ground color light brown. Genitalia 
(Fig. 7): with ostium bursae opening wide with sclerotized process cephalad; ductus sem- 
inalis joins ostium bursae at same point as ductus bursae; corpus bursae with two blunt 
siga; located far cephalad in abdomen, concurrent with segments A2—A4; papillae anales 
rounded, setose. 

Types. Holotype: male, Cerro Aguacatal, Mpio Riosucio, Caldas, Colombia, 15 April 
1993, leg. Salazar. Paratypes: 6 males, 2 females as follows: Cerro Aguacatal, Mpio Riosu- 
cio, Caldas 1300 m, 1 April 1994; Cerro Clavijo, Mpio Riosucio, 20 July 1994. The holo- 
type is deposited in the Museo de la Universidad Nacional, Bogota, Colombia. The 
paratypes are in the collections of C. Callaghan, J. LeCrom, and E. Schmidt-Mumm of 
Bogota, and the Universidad de Caldas, Manizales. 

Diagnosis. Calydna volcanicus is closely allied to Calydna hemis Schaus from south- 
eastern Brazil, and therefore is provisionally assigned to the same genus. However, both 
these species differ from Calydna thersander (Stoll), the type species of the genus, in the 
large, sexually dimorphic palpi and the structure of the genitalia, suggesting that they be- 
long to an undescribed genus. 

Distribution and Habits. The species was discovered by J. Escobar on two small 
mountain ranges, Cerro Aguacatal and Cerro Clavijo, both on the eastern slope of the 
Cordillera Occidental in the Municipio of Riosucio, Caldas Department between 1200 
and 1600 m above sealevel (Fig. 8). The climate is Very Humid Premontane Forest (IGAC 
1977) with a mean annual rainfall of 2000-4000 mm and a biotemperature of between 
18—24°C. The area has been extensively cultivated for coffee (Coffea arabica) with the 
consequent alteration of the original vegetation. The upper slopes of the Cerro Aguacatal 
(1300 m) are practically devoid of vegetation and a concrete cross is at the summit. Ca- 
lydna volcanicus flies below the summit in a deep canyon with low bushes growing on the 
slopes. Males perch on low bushes after 1200 h, resting with wings spread on the upper 
leaf surface. When disturbed, they fly rapidly, engaging other males in a whirling chase 
before returning to their original spot. The lower slopes of the Cerro Aguacatal 
(1500-1600 m) and the Cerro Clavijo are occupied with coffee plantations shaded by na- 
tive trees of the genera Inga and Albizzia. Here, males perch along trails through the 
plantations between 1300-1400 h, always on the upper leaf surfaces with wings spread. 
Females frequent the same areas and visit flowers. 

Etymology. The name volcanicus has no significance. 

Other riodinid species recorded from the same habitat include: Siseme pallas 


60 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 8. Map of Colombia. Hatched areas represent zones above 1000 m. Triangle in- 
dicates type locality of Calydna volcanicus. 


(Latreille), Catocyclotis elpinice (Godman), Amphiselenis chama (Staudinger), Melanis 
cratia (Hewitson), Adelotypa densemaculata (Hewitson), Charis nr. zama (Bates), and 
Calephelis schausi McAlpine. The discovery of Amphiselenis chama in north central 
Colombia constitutes an extension of its previously known range in the Venezuelan An- 
des. Examination of Amphiselenis from both Colombia and Venezuela revealed no consis- 
tent variation in phenotype between the two populations. 


ACKNOWLEDGMENTS 


We thank Luis Constantino and an anonymous reviewer for their helpful comments. 


VOLUME 51, NUMBER Il 61 


LITERATURE CITED 


Brown, K. B. 1979. Ecologia geografica e evolugao nas florestas neotropicais. Unpubl. 
Ph.D. Thesis, Univ. Estadual de Campinas, Sao Paulo, Brazil. 265 pp. 

CALLAGHAN, C. J. 1983. Notes on the genus Imelda (Riodinidae). J. Lepid. Soc. 37: 
254-256. 

HarVEY, D. H. 1987. The higher classification of the Riodinidae (Lepidoptera). Unpubl. 
Ph.D. Thesis, Univ. Texas, Austin, Texas. 266 pp. 

IGAC, 1977. Zonas de vida 0 formaciones vegetales de Colombia. Vol. xiii, no. 11. 238 pp. 

Kxots, A. B. 1970. Lepidoptera, pp. 97-111. In Tuxen, S. L. (ed.), Taxonomist’s glossary 
of genitalia in insects, 2nd revised and enlarged edition. Munksgaard, Copenhagen. 

MILLER, L. D. 1969 [1970]. Nomenclature of wing veins and cells. J. Res. Lepid. 8:37—48. 

SALAZAR ESCOBAR, J. A. 1993. Noticias sobre seis raras especies de licenidos colombianos. 
Descripcién de una nueva especie de Riodininae para Colombia (Lepidoptera: Ly- 
caenidae). Shilap 21(81):47—53. 

SALAZAR ESCOBAR, J. A. & L. M. CONSTANTINO. 1993. Descripcidn de cuatro nuevas es- 
pecies de Riodininae para Colombia (Lepidoptera: Lycaenidae). Shilap 21(81):13-18. 


Received for publication 10 January 1995; revised and accepted 7 January 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 62-74 


A REVISION OF THE EUSELASIA ORFITA COMPLEX 
(RIODINIDAE) 


CURTIS J. CALLAGHAN 
Avenida Suba 130-25, Casa 6, Bogota, Colombia 


ABSTRACT. The Euselasia orfita complex (Riodinidae) sensu Stichel 1919 is re- 
vised. Separate keys to adult males and females are presented, as well as notes on nomen- 
clature, geographical variation, distribution and adult habits. As revised, this complex in- 
cludes: E. orfita (Cramer 1777); E. eutychus (Hewitson 1856) reinstated status, =E. 
ferrugo (Bates 1868) new synonym; E. cuprea Lathy 1926 reinstated status; E. cyanira 
new species; E. clithra (Bates 1868), =E. clithra jugata Stichel 1919 new synonym; and 
E. phedica (Boisduval 1836). 


Additional key words: neotropical, South America, Brazil, Colombia, Peru, 
Ecuador. 


The purpose of this paper is to revise the Euselasia orfita complex 
(Riodinidae), a small assemblage of species inhabiting South America 
east of the Andes. Although common in museum collections, these spe- 
cies are difficult to identify because they are similar in appearance, es- 
pecially females. This revision is preliminary because other Euselasia 
species may belong to the E. orfita group. For this reason, a phylogeny 
will be attempted only later, when confidence about the systematics of 
this large and varied genus is on a firmer foundation. 


MATERIALS AND METHODS 


In addition to the author's collection (CJC), the following institutional 
collections were consulted during this study: Museu Nacional, Rio de 
Janeiro (MN); Smithsonian Institution, Washington D.C. (NMNH); Mu- 
seum National d’Histoire Naturelle, Paris (MNHN). A total of 256 spec- 
imens was examined, including 36 specimens taken on loan. Thirty 
eight genitalic preparations were made by soaking the abdomens over- 
night in 10% potassium hydroxide solution; genitalia were stored in vials 
cross referenced to the specimens. Terminology of the genitalic struc- 
tures follows Klots (1970), and the designation of wing veins and cells 
follows Miller (1969). Measurements were made using a binocular mi- 
croscope fitted with an ocular micrometer. Field observations were 
made by CJC unless otherwise indicated. 


MORPHOLOGY OF THE EUSELASIA ORFITA COMPLEX 


The species of the E. orfita complex are easily separated from other 
Euselasia by their large size (forewing length 17-24 mm), and the series 
of parallel dark red/brown bands that cross the ventral surface of both 
wings. Other distinguishing characteristics of the complex are as fol- 
lows: 


VOLUME 51, NUMBER 1 63 


Fics. 1-8. Adults of Euselasia orfita complex. 1, E. orfita, dorsal, male (Brazil); 2, 
ventral, male, showing numbering of bands; 3, dorsal, female (Brazil); 4, ventral, female; 
5, E. cuprea, dorsal, male; 6, ventral, male; 7, dorsal, female; 8, ventral, female; 


Dorsal wing pattern. The male dorsal wing pattern characterizing the E. orfita group 
can best be seen using light projecting from behind the viewer's shoulder, with specimens 
tilted away from the viewer from the apex towards the base of the wings. There are three 
distinct arrangements of irridescent blue scaling on the forewing. In the first, the color is 


deep purple and confined to the basal half (Fig. 1). In the second, the color is lighter blue 


64 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


and the pattern consists of a wide >3 mm band beginning at the costa above the cell, that 
curves around the end of the cell and terminates at the inner margin, and with darker 
blue-violet between the band and the base; the distal area lacks iredescent scaling (F ig. 
9). In the third, there is a thin band 1 mm wide beginning at the base that continues along 
the margin to distad of the cell, and then crosses the wing as a 2 mm wide band to the in- 
ner margin, with dark blue scaling on both forewing and hindwing separated from the 
thin band by a black area (Fig. 21). 

The pattern on the hindwing likewise consists of three types: a deep purple tone in the 
distal half (Fig. 1); a marginal band of light blue scales (Fig. 9); or a white limbal area with 
a glossy light blue sheen (Fig. 25). 

Ventral wing pattern. The E. orfita group is characterized by a ventral forewing pat- 
tern that includes six bands crossing the forewing from the costal margin to 2A, and the 
hindwing from the costa to the inner margin. These are numbered 1-6 from the base to 
the submargin in Fig. 2. Bands 1-4 are reddish brown and band 1 is thinner and shorter. 
The first three bands are slightly convergent toward the forewing costa. Band 5 is broken 
into a series of figures consisting of spots and lines, and band 6 follows the margin of both 
wings as a distinct band or shading. The bands are the principal means of associating the 
Sexes. 

On the forewing, the shape of band 4 may be straight (Fig. 10) or S-shaped (Fig. 6) in 
both sexes. Band 5 begins as three arrow-shaped spots in cells R,—R,, Ry-M,, and M,—Mg, 
followed by a wide band between M, and 2A on the inner margin. The band between M, 
and 2A has three types: a straight heavy line to 2A, veering basad to the inner margin as a 
thin projection (Fig. 2); a band curved towards the costa, uniting with band 4 at M, (Fig. 
6); straight as above, but not reaching below 2A (Fig. 10). 

The ventral hindwing is characterized by the continuation of bands 1-4 from the 
forewing. Band 5 becomes a series of arrowhead-shaped spots pointing basad between 
the veins, with a blue ocellus in cell M,—M;, bordered on three sides with orange and dis- 
tad by white. The distance between the ocellus of the hindwing ventral surface and band 
3 differs significantly between species, and is greater in females. 

Genitalia. The male genitalia (Figs. 29-34) are simple with a broad bilobed uncus 
widely separated from the tegumen, the latter with two narrow falces and joined to the 
valvae by a long, thin vinculum. The saccus consists of a widening of the base of the vin- 
culum. The valvae are elongate, spatula-shaped with flat tips and a heavily sclerotized pro- 
cess near the base. The aedeagus is blunt or pointed, with a coiled vesica and is guided by 
a sclerotized triangular-shaped transtilla. There is considerable intraspecific variation. 

In the female genitalia (Figs. 35-39) the lamella postvaginalis is bladelike with a small 
point between the two blades. The posterior ductus bursae has two tiny crescent-shaped 
signa. There appeared to be considerable differences among the genitalia examined; how- 
ever, the number of preparations was insufficient to separate specific from individual vari- 
ation with confidence. 

Frons and palpi. The lateral margins of the frons and palpi are either dirty yellow or 
white. 

Geographic distribution. The E. orfita complex is limited to tropical South America 
east of the Andes from the Guianas across the Amazon basin to Peru (Bolivia?). In the 
Andes the complex occurs up to 1000 m (Fig. 40). 


PROPOSED CLASSIFICATION 


In his revision of Euselasia, Stichel (1928) recognized nine species 
in this complex (Orfitoformes): E. orfita (Cramer), E. ferrugo (Bates), 
E. clithra (Bates), E. phedica (Boisduval), E. eurymachus (Hewitson), 
E. eurysthenes (Hewitson), E. orba Stichel, E. issoria (Hewitson) and 
E. euodia (Hewitson). Examination of the latter five species suggests 
that they are not related to the Euselasia orfita complex. They lack the 
six band pattern on the wing ventral surface typical of E. orfita. On 


VOLUME 51, NUMBER 1 65 


13 


14 


15 


16 


Fics. 9-16. Adults of Euselasia orfita complex. 9, E. eutychus, dorsal, male (Brazil, 
AM); 10, ventral, male; 11, dorsal, female (Brazil, AM); 12, ventral, female; 13, dorsal, 
male (Brazil, PA); 14, ventral, male; 15, dorsal, female (Brazil, PA); 16, ventral, female. 


E. eurymachus and E. eurysthenes the tips of the valvae are bifurcated 
with the lower projection longer and turned inwards, a characteristic of 
the Euselasia anica complex (Aniciformes sensu Stichel). Although the 
male genitalia of E. orba, E. euodia and E. issoria are close to E. orfita, 


66 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 17-20. Adults of Euselasia orfita complex. 17, E. eutychus, female (Brazil, PA); 
18, ventral, female; 19, E. cyanira, holotype, dorsal; 20, ventral. 


the wing pattern suggests that they form a distinct monophyletic group. 
Thus, all five taxa are omitted from consideration in this paper. The fol- 
lowing classification for the E. orfita complex is proposed: 


E. orfita (Cramer 1777) 
=E. orfita eutychus f. truculenta Stichel 1924 
E. eutychus (Hewitson 1856), reinstated status — 
=E. ferrugo (Bates 1868), new synonym 
=E. dyrrhachius Seitz 1913 
=E. eutychus f.lacteata Stichel 1919 
=E. orba spectralis f. pallida Lathy 1926 
E. cyanira, new species 
E. cuprea Lathy 1926, reinstated status 
E. clithra (Bates 1868) 
=E. jugata Stichel 1919, new synonym 
E. phedica (Boisduval 1836) 


KEY TO MALES OF THE E. ORFITA COMPLEX © 


la. Dorsal forewing with a blue band across forewing from end of cell to inner 

Naf g2 10 earn Re Ce EA NIM a revel ai enc Menu E ORME aS) cuOMu NaI hig 8 ec co oe ass 4 
1b. Dorsal forewing dark purple with no transverse band................-.---- D 
2a. On ventral surface of forewing, space between lines three and four lighter than 


VOLUME 51, NUMBER 1 67 


Fics. 21-28. Adults of Euselasia orfita complex. 21, E. clithra, dorsal, male (Brazil, 
PA); 22, ventral, male; 23, dorsal, female (Brazil, PA); 24, ventral, female; 25, E. phedica, 
male (Fr. Guiana); 26, ventral, male; 27, dorsal, female (Fr. Guiana); 28, ventral, female. 


ground color; line four straight, line five to CU,; distance between ocellus 

and third band 0.8—1.0 mm; valvae short, flat, tips rounded.............. E. orfita 
2b. On ventral surface of forewing, ground color uniform; line four S-shaped; line five 

Gilg camnomi2 nv tovNiealineshive touMl, siccue) iil. sa 8) eG en bee Lies 3 


68 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


3a. Hindwing dorsal surface black with dark blue margin; ocellus distance 

0.7—1.0 mm; valvae wide, tips flat, spatula-shaped.................... E. cuprea 
3b. Hindwing dorsal surface with white limbal area; line three on ventral forewing 

S-shaped; line four curved and yellow; line five reaches M3; ocellus distance 

0.3—0.8 mm; valvae very short, slightly flattened.................... E. phedica 
4a. Forewing with wide >3 mm blue band crossing discal area to inner margin... . . 5 
4b. Forewing with thin <3 mm blue band from base to end of cell, then crossing 

wing to tornus; ventral forewing line five wide to 2A, then turning basad as 

a thin line to inner margin; populations in eastern Amazon basin with bands 

three and four more widely separated with white scaling; ocellus distance 

0.8—1.8 mm; valvae long, round, slightly curved inwards, tips rounded. . . . E. clithra 
5a. Facial sutures white; dorsal wing surface with ventral surface markings 

showing through; ventral surface covered uniformly with a blue glaze; line 

four reaching beyond 2A as thin line turning basad to inner margin; 

ocellus distance 0.7—1.1 mm; valvae like E. clithra, slightly longer ...... E. cyanira 
5b. Facial sutures yellow; dorsal wing surface opaque; ventral surface blue 

glaze confined to base of forewing and anal angle of hindwing; line four 

stopping at 2A; ocellus distance 0.5—0.9 mm; valvae very long, narrow, tips 

pointed: and: turned imwards 22.4... 6. Pe ee, ee ee E. eutychus 


KEY TO FEMALES OF THE E. ORFITA COMPLEX 


la. Ventral forewing lines four and five curved costad, meeting at Mj............. 2 

Ib:-Ventral forewing lines four and! five straight. 92) 4. 2 3 

2a. Ventral hindwing ocellus in cell M,—M, round, less than 0.9 mm from band four; 
band four on forewing yellow; dorsal grey-blue; ocellus distance 0.96 mm 


Pe ae ioe manag Oe tS mer MaRS cas ie eg bo! co a 3 E. phedica 
2b. Ventral hindwing ocellus in cell M,—M, <1.1 mm from band three; band four on 
forewing reddish brown; dorsal brown; ocellus distance 1.1mm........ E. cuprea 


3a. Palpi, facial sutures yellow; ocellus on ventral hindwing <1 mm from band four; 

dorsal hindwing ocellus arrowhead-shaped; individuals may have white _ 

discal area ventrad or dorsad; ocellus distance 0.7-1.0mm...........E. eutychus 
3b. Palpi, facial sutures white; ocellus on ventral hindwing <1 mm from band four... 4 
4a. Dorsal hindwing with yellow scaling around ocellus; ventrally with ground 

color between bands three and four slightly lighter; forewing lines five and 

six wide, broken, indistinct; ocellus distance 1.2-1.4mm............... E. orfita 
4b. Dorsal hindwing without yellow scaling; ventral forewing bands five and six 

narrow, of uniform width, unbroken; ground color uniform; ocellus distance 

HG OV nana eo ac nN NT 2 Meare as nema ei at E. clithra 


SPECIES ACCOUNTS 


Euselasia orfita (Cramer 1777) 
(Figs. 1-4, 29, 35) 


Nomenclature. Cramer (1777) described Euselasia orfita from a male from Surinam. 
Figures D and E on his plate 112 are crude, but recognizable as E. orfita. The type appar- 
ently has been lost, and there is no specimen at Leiden that could be designated as a lec- 
totype. However, the species is distinct. 

Geographical Variation. The species shows little variation over its range from the 
Guianas to Brazil (Para, Amazonas). A single female from “Santa Cruz Bolivia” at the 
MNHN is probably mislabeled. 

Ecology and Behavior. Euselasia orfita inhabits terra firme forests in the lowlands of 
the Amazon basin and the Guianas. Males are encountered rarely in the forests perching 
in the early afternoon in treefalls and other small clearings, resting under leaves near the 
ground with their wings closed. When searching for oviposition sites, female E. orfita of- 
ten rest on the upper surfaces of leaves, and this habit, along with the barred wing under- 
sides and loping flight, make it easy to confuse them with satyrids. 


VOLUME 51, NUMBER Il 69 


SVE 
\ 
y 


i | 


Fics. 29-34. Male genitalia of Euselasia orfita complex. 29, E. orfita; 30, E. cuprea; 
31, E. eutychus; 32, E. cyanira; 33, E. clithra; 34, E. phedica. 


Material Examined. FRENCH GUIANA: St. Elie pk 15.5 on D21, 76 (NMNH); 
Moyen Maroni, ld (MNHN); Kana (Mana?), Id (CJC); St. Laurent, 4d 12 (MNHN). 
BRAZIL: Obidos, Pa, ld (NMNH), 36 (MNHN); Trombetas, Pa., 1d (MNHN); Tapajés, 
Pa, ld (MNHN); Manaus, Am, I2 (MNHN); Tefé, Am, 12 (MNHN); 80 km N. Manaus, 
Am, 7¢ 4° (CJC). BOLIVIA: Santa Cruz, 12 (MNHN)(?). 


Euselasia cuprea Lathy 1926, reinstated status 
(Figs. 5-8, 30, 36) 


Nomenclature. This species was described from a male, currently in the British Mu- 
seum (Natural History). The type locality is St. Laurent, Maroni River, French Guiana. 
Stichel (1928) treated E. cuprea a subspecies of E. orfita. However, the two are distinct 
morphologically, and sympatric. 

Geographical Variation. The species is confined to the Guianas and shows no varia- 
tion among the material examined. 


70 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 35-39. Female genitalia of Euselasia orfita complex. 35, E. orfita; 36, E. 
cuprea; 37, E. eutychus; 38, E. clithra; 39, E. phedica. 


Ecology and Behavior: The species inhabits the terra firme forests of French Guiana. 
I have no personal experience with its habits. 

Material Examined. FRENCH GUIANA: Rte. de l'Est, ld (NMNH); St. Jean Ma- 
roni, lS (NMNH); PK43.5 N2, 3d (CJC); Nouveau Chantier, 26 22 (MNHN); “Guyane 
Frangaise,” 25 (MNHN); T. Laurent, I2 (MNHN); St. Domenti, 1d (CJC); Galion, 1d 
(CJC). 


Euselasia eutychus (Hewitson 1856), reinstated status 
(Figs. 9-18, 31, 37) 


=Euselasia eutychus f. pallida Lathy 1926 
=Eurygona ferrugo (Bates 1868) 


VOLUME 51, NUMBER 1 yal 


8 
5 I> 
“ D_A @&. 
@.~ A 
Oss AO 
O WO 
O 
A 
(eo) 
A 
A 
@ 


40 


Fic. 40. Geographical distribution of the Euselasia orfita complex. Filled circles = E. 
orfita, filled triangles = E. eutychus, filled squares = E. phedica, open circles = E. clithra, 
open triangles = E. cyanira, open squares = E. cuprea. 


=Euselasia dyrrachius Seitz 1913 
=Euselasia orba spectralis f. lacteata Stichel 1919 
Nomenclature. Hewitson (1856) described E. eutychus from a male, presently in the 
British Museum (Natural History), with the label indicating “Amazon” as the locality. His 
figures 44 and 46 are good representations of the species. Stichel (1919) placed E. euty- 
chus as a subspecies of E. orfita with no explanation. As in the case of E. cuprea, this was 
in error, as the two are distinct morphologically, as well as partially sympatric. 
Geographical Variation. There is significant variation both within and between pop- 
ulations of E. eutychus from Parad and Amazonas, Brazil, south of the Amazon River, to 
the foothills of the Andes from Meta, Colombia, south to Peru. In males from Para, band 
5 converges sharply from the anal angle towards band 4; in males from Amazonas, Brazil, 
Peru and Colombia this band is parallel to band 4. In some males, bands 3 and 4 fuse 


72 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


together (ferrugo Bates). Females show variability paralleling that of the males. Those 
from Para are generally darker than those to the east and have wider bands. Some indi- 
viduals have white ventral hindwing surfaces, and others have white on the discal area of 
the dorsal surface (pallida Lathy, and lacteata Stichel; Figs. 17-18). The white scaling is 
variable and is rare although present in all E. eutychus populations. 

Ecology and Behavior. Euselasia eutychus is widespread, from Para, Brazil, through- 
out the Amazon Basin to Peru. Unlike other members of the group, it adapts to disturbed 
habitats where it may be locally common. I have found the males perching in the late 
morning inside the edges of large clearings, resting on ventral leaf surfaces with their 
wings folded. Females are found less commonly within the forests. 

Material Examined. BRAZIL: Jaru, RO, 3d (CJC); Cuiaba-Santarem km 1666, Pa, 4d 
(CJC), ld 12 (NMNH); Vila Bela, MT, ld (CJC); Vilhena, RO km23, 26 (CJC); Manaus, 
AM, Id ( (CJC); Trombetas, Pa, 25 12 (MNHN); Faro, Pa, 26 12 (MNHN); Maués, Am, Id 
(MNHN); Tefé, Am, ld (MNHN); Conceigao, Tapajés, Pa, 35 (MNHN); Santarém, Pa, 4d 
(MNHN); Tapajés, Pa, 25 (MNHN); Itaituba, Pa, lI (MNHN); Amazon Sup., 26 1° 
(MNHN); Vista eae Rio Jurua Mirim, Acre, ld (KB). COLOMBIA: Puerto Inirida, Pu- 
tumayo, Id (CJC); La Macarena, Meta, ne ( (CJC); Leticia, Am, 2¢ (CJC); Villagarzon, Ca- 
queta, 3d; Rio Negro, Meta, 5d (CJC), 66 (NMNH); Willavicencies Meta, Id (CJC), 3d 29 
(MNHN); Montanita, Caqueta, 1d (CJC), 25 (NMNH); Tres Esquinas, Caqueta, 3d 29 
(NMNH); Putumayo, 85 (MNHN); Umbria, 1d (MNHN); Meta, nr. Villavicencio, 1d 
(NMNH). ECUADOR: Rio J’ondachi, 1d (CJC); Puyo Pastaza 26 (CJC), 2¢ (NMNH); 
Cotundo Napo, 2¢ (CJC), 4d 19 (NMNH), 26 (MNHN); Limoncocha 36 (NMNH). 
PERU: Tingo Maria, 3d (CJC); Chanchamayo, 26 (NMNH); Iquitos, 25 (NMNH), I? 
(MNHN), 6d (MNHN); Chanchamayo, 46 2° (MNHN); La Merced, 26 22 (MNHN); Rio 
Tono, 4d 29 (MNHN); Jepelacio, 1d (MNHN); Rio Perene, 35 (MNHN); Huancabamba, 
16 (MNHN); Madre als Dios, I (MNHN). 


Euselasia cyanira Callaghan, new species 
(Figs. 19-20, 32) 


Description. Male: legs yellow, palpi and facial sutures white; forewing lenath of holo- 
type 22 mm (range of material examined 20-22 mm, n=7); dorsal surface of forewing 
ground color black, costal margin purple-blue from base to end of cell, continuing as a 4 
mm wide lighter blue band to anal angle; hindwing ground color black, costa light brown, 
2 mm wide marginal blue band from M, to anal angle; ventral pattern shows through to 
dorsal surface; ventral surface light grey with a light blue sheen, stronger at the anal an- 
gle; on forewing, band | is absent; bands 2—4 red/brown and parallel; band 5 broken into 
three arrow shaped spots in cells R;—R,, R,-M, and M,—Mg, continuing as a wide band 
from M, to 2A, then as a thin line at an angle basad to the inner margin; band 6 distinct, 
reaching CU,; margin light brown with black fringe; hindwing bands 2—4 are continua- 
tions of those on the forewing from costa to 2A; bands 2 and 3 slightly concave to the 
base; band 5 consists of elongated arrow-shaped spots pointing basad, two each in cells 
CU,—CU, and CU,-2A and the rest with one each, except cell M,—M, which contains an 
oval, black ocellus separated from line 3 by a yellow shading 0.87 mm wide and bordered 
distad by a short, white line; margin light grey with a 1 mm wide orange line distad, and a 
thin black line at base of fringe. Female: unknown. 

Types. Holotype: male, with label “Pumayacu, Huallaga, Peru,” and a red TYPE label. 
Paratypes: six males from Manaus, Amazonas and U. Putumayo, SE Colombia. The holo- 
type and two paratypes are in the Museum d Histoire Naturelle, Paris; one paratype is in 
the collection of the author; material will be distributed to the Museo Nacional, Rio de 
Janeiro and the USNM, Washington, D.C. 

Genitalia. As illncerited (Fig. 32). 

Geographical Variation. Euselasia cyanira ranges from Manaus, Brasil to Putumayo, 
Colombia then to central Peru, always in humid tropical lowland forest All specimens ex- 
amined are similar in appearance, suggesting that the species is quite uniform over its 
range. 

Dingess The dorsal surface of E. cyanira resembles closely that of E. eutychus with 


VOLUME 51, NUMBER 1 1} 


broader blue bands on the forewings. However, E. cyanira may be easily separated by its 
white rather than yellow facial sutures. Although the ventral surface of E. cyanira resem- 
bles that of E. clithra, hindwing bands 2 and 3 are slightly concave to the base in E. 
cyanira and straight in E. clithra. The wider blue band on the forewing of E. cyanira sep- 
arates it from E. clithra. 


Euselasia clithra (Bates 1868) 
(Figs. 21-24, 33, 38) 


=E. clithra jugata Stichel 1919, new synon 

Nomenclature. Euselasia clithra was described from a male captured in “Para, L. 
Amazonas” by H. W. Bates. The type specimen is in the British Museum (Natural His- 
tory). 

Pa eiaphical Variation. Euselasia clithra shows considerable clinal variation over its 
range from eastern Para to the foothills of the Andes south of the Amazon and Rio Negro 
rivers. The males from Para, nominate E. clithra, are characterized by ventral hindwing 
bands 2 and 3 diverging from band 4 toward the inner margin, leaving a wide, open space 
of white scales between them. To the west, the lines become increasingly closer together 
until they are nearly parallel (jugata Stichel 1919, described from Rio Jurua, Brazil). The 
dorsal surface invariable. Females tend to be slightly darker in the western portion of the 
range. 

Pelee and Behavior. In my experience, E. clithra inhabits the terra firme forest 
where the males perch at the edges of treefalls and small clearings in the late morning and 
early afternoon hours. Adults rest near the ground under sunlit leaves with wings closed. 
I have never found them common, especially the females, but judging from the long se- 
ries in museums, at times they may be encountered more frequently. 

Material Examined. BRAZIL: Cuiabaé-Santarem km 1666, Pa, 2d (CJC), 16 (NMNH); 
Ariquemes, RO, Id (CJC); Manicoré, Am, ld (CJC), 25 (MNHN); Tapajés, Pa, 1d 
(MNHN): Monte Cristo, Pa, lé (MNHN); Itaituba, Pa, lé (MNHN); Altamira, Pa, 1d 
(MNHN); Barreiras, Pa, 4d (MNHN); Conceicao, Pa, 86 22 (MNHN); Amazonas, 1d 
(MNHN); Massauary, Rio Negro, Am, ld (MNHN); Uypiranga, Am, 26 (MNHN); Sao 
Paulo de Olivenga, Am, Id 19 (MNHN); Rio Como?, If? (MNHN); Manacapuru Rd., south 
of Manaus, Am, Id (CJC). PERU: Pakitza, 206 3° (NMNH); Iquitos, 4d (MNHN); Ama- 
zon Supl., ld (MNHN); COLOMBIA: Putumayo, 66 12 (MNHN); Umbria, Putumayo, 1° 
(MNHN). 


Euselasia phedica (Boisduval 1836) 
(Figs. 25-28, 34, 39) 


Nomenclature. Boisduval named E. phedica from a male illustrated from French 
Guiana. The type is in the British Museum (Natural History). 

Geographical Variation. There is no variation over the range of this species from 
French Guiana through the Amazon drainage north of the Amazon river to southern 
Venezuela. A male in the MNHN collection from “Putumayo” may be mislabeled. 

Ecology and Behavior. [ have no personal experience with this butterfly. However, 
K. Brown (pers. comm.) says that it inhabits deep primary forest, perching close to the 
ground under leaves on the edge of treefalls. 

Material Examined. FRENCH GUIANA: Bas Maroni, 66 (MNHN); St. Laurent, 1¢ 
(MNHN); St. Elie, pk 15.5 on D21, 26 (NMNH); St. Jean, Maroni, 2d (NMNH). 
VENEZUELA: Cerro de Neblina, Basecamp, 140 m, 12 (NMNH). BRAZIL: Ypiranga, 
Am, ld 1° in cop. (MNHN); Manaus, Am, ld (MNHN); Obidos, Pa, 1d (MNHN); Rio 
Umary, 4d (MNHN). COLOMBIA: Putumayo, Id (MNHN)(?). 


ACKNOWLEDGMENTS 


I am indebted to the curators of the museums visited, especially R. Robbins and M. 
Jacques Pierre at the Smithsoninan and Museum d Histoire Naturelle, respectively, for 


74 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


access to their collections for the examination and loan of material: and to R. Robbins for 
his orientation during the preparatory phase of the project. To Keith Brown of the Uni- 
versidade Estadual de Campinas, Sao Paulo, Brazil, my thanks for the loan of much valu- 
able material and comments on the manuscript. I also thank Don Harvey of the Smith- 
sonian for many helpful comments on the manuscript. 


LITERATURE CITED 


BOISDUVAL, J. B. 1836. Spécies général des lépidoptérés. Paris, Roret. 

BATES, H. W. 1868. A catalogue of Erycinidae, a family of diurnal Lepidoptera. J. Linn. 
Soc. Lond., Zool. 9:367—459. 

CRAMER, P. 1777. De Uitlandische Kapellen voorkomende in de drie Waereld-deelen 
Asia, Africa en America, Amsterdam, Baalde 2. 151 pp. 

HEwITSON, W. C. 1856. Illustrations of new species of exotic butterflies, selected chiefly 
from the collections of W. Wilson Saunders and William C. Hewitson. London. Van 
Voorst. L(18): p.[54]. 

KiotTs, A. B. 1970. Lepidoptera, pp. 115-130. In Tuxen, S. L. (ed.), Taxonomist’s glossary 
of genitalia in insects, 2nd revised and enlarged edition. Munksgaard, Copenhagen. 

LaTny, P. 1926. New species and forms of the genus Euselasia (Lepidoptera) in the Joicey 
collection. Entomologist 59:143—146. 

MILLER, L. D. 1969. Nomenclature of wing veins and cells. J. Res. Lepid. 8:37—48. 

SEITZ, A. 1913. Die Grossschmetterlinge der Erde. Stuttgart, Kernen. 5: pl. 121. 

STICHEL, H. 1919. Vorarbeiten zu einer Revision der Riodinidae Grote IV. Deut. Ent. 
Zeit. 1919:161—171. 

. 1924. Beitrige zur Kenntnis der Riodinidenfauna Sudamerikas VII. Nord 

Brasilien (Amazonas). Zeitschrift Wiss. Ins. Biologie. 19:245—250. 

. 1928. Lepidoptera Nemeobiidae. Das Tierreich 51. xxx + 330 pp. 


Received for publication 1 September 1993; revised and accepted 7 January 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 75-82 


TERMINATION OF PUPAL DIAPAUSE IN CALLOPHRYS 
SHERIDANII (LYCAENIDAE) 


KIYOSHI HIRUMA 


Department of Zoology, University of Washington, Box 351800, Seattle, 
Washington 98195, USA 


JONATHAN P. PELHAM 


Burke Memorial Museum, University of Washington, Box 353010, Seattle, 
Washington 98195, USA 


AND 


HERVE BOUHIN 


URA CNRS 674, Laboratoire de Zoologie, Université de Bourgogne, 
F-21000 Dijon, France 


ABSTRACT. When Callophrys sheridanii neoperplexa/newcomeri pupae were incu- - 
bated at 4°C for 100 days, 65% of the pupae eclosed within 8 days and the remaining 35% 
eclosed gradually 45 to 200 days after the termination of the chilling. Non-chilled pupae 
stopped their development at the eye pigmentation stage. More than 60% of adult eclo- 
sion was observed within 30 min after lights on. These results suggest that adult eclosion 
of this species occurs abruptly in the early morning on the first few warm days in spring, 
and one of the factors explaining the sporadic late records in the field is the gradual ter- 
mination of the pupal diapause. 


Additional key words: circadian rhythm, eclosion, low temperature, endocrinology. 


Insect development is governed primarily by ecdysteroids and juve- 
nile hormone (JH). Prothoracicotropic hormone (PTTH) from brain 
neurosecretory cells stimulates prothoracic glands to secrete 3-dehy- 
droecdysone in Manduca sexta L. (Sphingidae) (Warren et al. 1988), it 
is then converted to ecdysone and then to an active hormone, 20-hy- 
droxyecdysone, which initiates development (Rees 1989). Ecdysteroids 
initiate the larval molt in the presence of JH secreted from corpora al- 
lata. In the last instar larva in Lepidoptera, a small peak of ecdysteroids 
in the absence of JH causes the cessation of feeding, larval-pupal com- 
mitment of the epidermis, and the onset of wandering behavior. Subse- 
quent exposure to ecdysteroids initiates pupation in the presence of JH 
(Riddiford & Hiruma 1990). This JH in the wandering stage is impor- 
tant to coordinate PTTH release (Hiruma 1986) and subsequently it 
stimulates prothoracic glands to secrete ecdysteroids that are responsi- 
ble for pupation, in addition it prevents to become an adultoid (a pupa 
with some adult structures) in some lepidopteran species (see Hiruma 
1980). Adult development is caused by an ecdysteroid surge in the pu- 
pal stage in the absence of JH (see Riddiford & Hiruma 1990 and Nij- 
hout 1994 for reviews). 


76 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Pupal diapause is characterized by the failure of the ecdysteroid se- 
cretion from prothoracic glands after pupation, which is primarily due 
to the failure of the release of PTTH from brains (Williams 1946, 1947, 
Denlinger 1985). The brain of a diapausing pupa can be activated to se- 
crete PTTH by chilling, thus terminating diapause so that adult devel- 
opment is initiated (Williams 1952). 

Callophrys sheridanii Edwards (Lycaenidae) is widely distributed in 
the western U.S., and differentiated to several different subspecies (Scott 
1986). In Washington State, there are two subspecies distinguished: C. s. 
neoperplexa Barnes and Benjamin occurring in the Columbia Basin and 
adjoining regions, and C. s. newcomeri Clench occurring in the south, 
southeast, and the east slope of the Cascade Mountains. The two inter- 
grade in the east central Cascades, from lowland steppe to high moun- 
tains in Kittitas and Chelan Counties west of the Columbia River (Pel- 
ham & Hiruma, pers. obs.). This species is considered univoltine and 
enters obligatory diapause in the pupal stage for hibernation (Scott 1986). 

In this paper, we show that diapause intensity in C. s. neoperplexa/ 
newcomeri pupae is heterogeneous and the timing of the adult eclosion 
responds to photoperiod. Also, we discuss the relationship between the 
results obtained in the laboratory and those of field observations. 


MATERIALS AND METHODS 


Eggs and first instar larvae of Callophrys sheridanii neoperplexa/new- 
comeri were collected from Eriogonum compositum Dougl. (var. 
leianthum Hook.) (Polygonaceae) in Schnebly Coulee, Kittitas County, 
Washington, and some of the eggs were laid in the laboratory on E. 
compositum by females caught in the same location. All the field caught 
eggs and larvae were brought into the laboratory, and reared under 
crowded conditions on E. compositum leaves in a plastic dish (14 cm di- 
ameter/2 cm height) at 26°C in a 17L:7D photoperiod. Twenty to 30 lar- 
vae were reared together until third instar larvae, then reduced the 
numbers to 10 to 15 in the fourth instar larvae. The leaves were changed 
daily. The food plant was wrapped with plastic bags and kept at 4°C no 
longer than 2 weeks. Lights off, the beginning of a new day, was set at 
00:00 AZT (Arbitrary Zeitgeber Time) (Pittendrigh 1965). In this condi- 
tion, cannibalism in larvae of this species was not observed as observed 
in the closely related European species, Callophrys rubi L. (Lycae- 
nidae) (Ford 1945). 

Pupae were kept at 26°C in a 17L:7D photoperiod for 67 to 77 days 
after pupation, they were then transferred to a 4°C incubator for 100 
days at dark, followed by transferring back to the 26°C condition 
(17L:7D). Non-chilled pupae were kept at 26°C at dark for 100 days in- 
stead of placing at 4°C, then at 26°C in a 17L:7D condition. Adult eclo- 


VOLUME 51, NUMBER 1 WL 


25 
male : 
20 — Non-chilled 
C7 female 5 
0 el 
150 200 
50 100 150 200 


Days after chilling 


Fic. 1. Adult eclosion from Callophrys sheridanii neoperplexa/newcomeri pupae. Pu- - 
pae were kept at 26°C for 67 to 77 days after pupation, they were then transferred to 26°C 
condition 100 days after the incubation at 4°C (n = 122). One male eclosed and 2 pupae 
died at 4°C. Nineteen pupae died after the chilling, but 15 developed to adult inside of 
the pupal cuticles and died without eclosion, probably because of desiccation. As a con- 
trol, pupae (n = 50) were kept at 26°C throughout the experiments (see Materials and 
Methods), and the days of eclosion were calculated as 0 at the time of the termination of 
the chilling in the experimental (inset). Only 6 eclosed, and one developed to adult but 
did not emerge (“150 days” is equivalent to “327 days after pupation”). The rest of them 
died by desiccation without forming any scales. Dead pupae were excluded from the data. 


sion was checked daily for about 400 days after pupation, and dissected 
all the non-eclosed pupae to examine whether or not they died. 


RESULTS 


Time of eclosion of diapausing pupae. All pupae developed to the 
stage to which compound eye pigmentation occurred within 67 to 77 
days at 26°C in a 17L:7D photoperiod, then stopped their development 
(n = 202). Most of the pupae produced very audible creaking sounds 
when disturbed as reported in many lycaenid pupae (Downey 1966, 
Brakefield et al. 1992), which was predicted by the presence of the 
stridulating organ in C. sheridanii (Downey 1966). The sound was pro- 
duced from shortly after pupation to the beginning of the eye pigmen- 
tation stage. 

Figure 1 shows that if pupae were not exposed to 4°C, only 12% of 
the pupae eclosed (6 out of 50) 326 to 377 days after pupation, which 
were equal to 149 to 200 days calculated as days after chilling (see be- 
low). When pupae were placed at 4°C for 100 days, the adult eclosion 


78 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


occurred in 65% of the pupae within 8 days after the termination of the 
chilling. One male eclosed even at 4°C. The remaining 35% eclosed 
gradually over about 200 days after chilling (Fig. 1). Apparently, the pu- 
pal diapause was terminated by the incubation at 4°C, and the diapause 
intensity of this population is not homogenous. This pattern of adult 
emergency indicates that there are at least two physiologically different 
pupae in the population. One is very sensitive to a low temperature to 
break diapause and adult eclosion occurs within 10 days after the termi- 
nation of chilling (group 1), and the other is much less sensitive to a low 
temperature whose adult eclosion occurs 50 to 100 days after the termi- 
nation of chilling (group 2). Since non-chilled pupae did not emerge 
more than 140 days (Fig. 1), the adult eclosion of the group 2 pupae is 
considered to have responded to the chilling. 

Sex ratio. Sixty-four percent (42 of 66) of adults were male in the 
group 1, but only 39% (16 of 41) in the other groups (Fig. 1). Male ratio 
was Statistically higher in group 1 (P « 0.001), but female ratio was sta- 
tistically higher in the others (P < 0.003). The overall sex ratio (male/ 
male + female) of the adults in the laboratory reared population was 
0.542 and it was not significantly different at P > 0.05 (n = 107). Accord- 
ing to the records deposited in the Burke Memorial Museum of the Uni- 
versity of Washington and those of our private collection, males were 
more commonly caught than females in the gulch bottom of Schnebly 
Coulee, where the food plant occurs, in mid March to mid April, and 
the sex ratio was 0.77 (n = 188) (significantly different at P « 0.001). 

Timing of adult eclosion. Of the adults that emerged 1 to 5 days af- 
ter chilling, more than 60% did so within 30 min after lights on (Fig. 2). 
This indicates that adult eclosion most likely occurs in the early morn- 


ing in the field. 


DISCUSSION 


The duration of diapause under well-defined environmental condi- 
tions usually is quite consistent for a given population, and the insect’s 
capacity to respond to environmental cues can also be directed by its ge- 
netic potential, sex, food plants and maternal history (Denlinger 1985, 
Pratt & Ballmer 1993). Diapause termination is linked directly to a spe- 
cific environmental cues, and the exposure to a low temperature was 
one of the main factors in C. s. neoperplexa/newcomeri (Fig. 1). Yet the 
trait is quite polymorphic for the termination as found in Hyalophora 
cecropia L. (Saturniidae) (Waldbauer & Sternburg 1973) whose bi- 
modality for the termination of the pupal diapause has a genetic basis. 
Figure 1 suggests that the population of C. s. neoperplexa/newcomeri is 
genetically heterogeneous, which also is evidenced by the mixed de- 
grees of the postmedian line. The similar bimodal adult emergence was 


VOLUME 51, NUMBER 1 79 


Lo , 


30 


Number 


10 


0 ZA | 
06:00 07:00 .08:00709:00" 10:00. 11 -(00 12:00 
Time (AZT) 


Fic. 2. Distribution of adult eclosion for C. s. neoperplexa/newcomeri under 17L:7D. 
Observations were performed on adults eclosed 1 to 5 days after the termination of chill- 
ing (n = 58). Seven adults eclosed during the dark. 


also observed in the laboratory reared Incisalia mossii mossii Edwards 
(Lycaenidae) (Hiruma, unpubl. data). 

The developmental stage of the arrest of diapausing pupae varies 
among species. In M. sexta (Bell et al. 1975, Bowen et al. 1984) and H. 
cecropia (Williams 1946), pupae enter diapause shortly after pupation 


80 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


due to the inactivation of the brains, whereas in Luehdorfia japonica 
Leech (Papilionidae), an univoltine species, the pupae complete almost 
all adult development by the beginning of winter. Yet adult eclosion 
does not occur until the following spring (Hidaka et al. 1971). It is in- 
teresting that pupal development occurs up to a certain level before the 
onset of diapause in C. s. neoperplexa/newcomeri. Probably, the brain- 
prothoracic gland axis is still active to secrete ecdysteroid even after pu- 
pation, so that the pupal development occurs until eye pigmentation 
stage. The determination of PTTH and/or ecdysteroid titer in the he- 
molymph is necessary to ascertain this hypothesis. 

Adult eclosion occurs at specific times of day in M. sexta, Antheraea 
pernyi Guérin-Ménéville (Saturniidae) and H. cecropia; it is controlled 
by a circadian clock and has been classified as a “gated” event (Truman 
1985). For the gated adult eclosion in M. sexta, the brain contains the 
clock that determines when eclosion hormone release will occur. The 
adult eclosion of C. s. neoperplexa/newcomeri also is thought to be con- 
trolled by a biological clock, and this may be due to the release of eclo- 
sion hormone shortly after lights on. 

According to our results reported in this paper, more than a half of 
the pupal population of C. s. neoperplexa/mewcomeri at a given location 
must emerge in the early morning on the first few warm days in Spring, 
followed by sporadic emergence throughout summer. Our field obser- 
vation partially supports this hypothesis. Adults of this species fly in 
early March to mid April in the lowland steppe such as Schnebly Coulee 
(500 m), but the peak of the flying period of fresh adults in each year is 
usually 7 to 10 days (Hiruma & Pelham, pers. obs.), which is supported 
by the outbreak of the eclosion of the diapausing pupae shortly after the 
termination of chilling (Fig. 1). In the higher elevations such as Chum- 
stick Mountains (1600-1700 m) in Chelan County, Washington, and 
Reecer Creek Road (1400 m) in Kittitas County, Washington, adults ap- 
pear in early April to early May (depending on the snow melt), but 
there are sporadic late records in both locations. In Schnebly Coulee 
and its vicinity, a relatively fresh female specimen was caught on 7 May 
1988, two males on 16 May 1987 and a male on 20 May 1984 (we did not 
find any adults in mid April in these years), and in the Chumstick 
Mountains, a fresh female on 16 July 1991 (Hiruma & Pelham, pers. 
obs.). Similar late records have been reported many times in C. rubi, 
and it is debatable whether or not these late records are due to second 
generations (see Ebert & Rennwald 1991 for detailed discussions). 
Based on our results, these late records are most likely due to the grad- 
ual eclosion depending on the diapause intensity, although we cannot 
rule out the possibility of second generations. 

Sexual differences in diapause response is common (see Denlinger 


VOLUME 51, NUMBER 1 81 


1985 for examples), and C. s. neoperplexa/newcomeri seems to exhibit 
this feature. Males were more common than females in the field, which 
may be due not only to collecting efficiency, but also to fewer eclosions 
of females in spring. It is well known that a certain percentage of dia- 
pausing pupae in some species of Anthocharis (Pieridae) and Atropha- 
neura (Papilionidae) do not emerge in the expected year after hiberna- 
tion, but emerge in the same season 2 to 3 years later (Kawazoé & 
Wakabayashi 1976). It has been reported as an extreme example in Eri- 
ogaster lanestris L. (Lasiocampidae) that the adult eclosion of 15 pupae 
occurred 14 years after pupation (Van-Nuvel 1976), although a 7-year- 
diapausing period was frequently observed in this species (South 1907, 
Van-Nuvel 1976). This seasonal adaptation has a number of advantages 
for unexpected unfavorable environments. In C. s. neoperplexa/new- 
comeri, the genetic program may not be completed to do so, and as a re- 
sult it causes gradual adult eclosion during the same year. 

These results indicate that results obtained in the laboratory are able 
to explain some of the unsolved observations in the filed, and therefore 
both types of research will help to analyze ambiguous results. 


ACKNOWLEDGMENTS 


We thank Lynn M. Riddiford and James W. Truman for the use of an insect rearing 
room. 


LITERATURE CITED 


BELL, R. A., C. G. RASUL & F. G. JOACHIM. 1975. Photoperiodic induction of the pupal 
diapause in the tobacco hornworm, Manduca sexta. J. Insect Physiol. 21:1471-1480. 

BOWEN, M. F., W. E. BOLLENBACHER & L. I. GILBERT. 1984. In vitro studies on the role 
of the brain and prothoracic glands in the pupal diapause of Manduca sexta. J. Exp. 
Biol. 108:9—24. 

BRAKEFIELD, P. M., T. G. SHREEVE & J. A. THOMAS. 1992. Avoidance, concealment, and 
defense, pp. 93-119. In Denies, R. L. H. (ed.), The ecology of butterflies in Britain. 
Oxford Univ. Press, Oxford. 

DENLINGER, D. L. 1985. Hormonal control of diapause, pp. 353-412. In Kerkut, G. A. & 
Gilbert. L. I. (eds.), Comprehensive insect physiology, biochemistry and pharmacol- 
ogy, vol. 8. Pergamon Press, Oxford. 

DOWNEY, J. C. 1966. Sound production in pupae of Lycaenidae. J. Lepid. Soc. 20:129-155. 

EBERT, G. & E. RENNWALD. 1991. Die Schmetterlinge Baden-Wiirttembergs. Band 2. 
Tagfalter II. Ulmer, Stuttgart. 535 pp. 

ForD, E. B. 1945. Butterflies. Collins, London. 368 pp. 

HIDAKA, T., Y. ISHIZUKA & Y. SAKAGAMI. 1971. Control of pupal diapause and adult dif- 
ferentiation in a univoltine papilionid butterfly, Lwehdorfia japonica. J. Insect Phys- 
iol. 17:197—203. 

HiIRUMA, K. 1980. Possible roles of juvenile hormone in the prepupal stage of Mamestra 
brassicae. Gen. Comp. Endocrinol. 41:392-399. 

. 1986. Regulation of prothoracicotropic hormone release by juvenile hormone in 
the penultimate and last instar larvae of Mamestra brassicae. Gen. Comp. En- 
docrinol. 63:201—211. 

KAWAZOE, A. & M. WAKABAYASHI. 1976. Colored illustrations of the butterflies of Japan. 
Ist ed. Hoikusha, Osaka. 422 pp. 

NiHoutT, H. F. 1994. Insect hormones. Princeton Univ. Press, New Jersey. 267 pp. 


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RIDDIFORD, L. M. & K. HrruMA. 1990. Hormonal control of sequential gene expression 
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TRUMAN, J. W. 1985. Hormonal control of ecdysis, pp. 413-440. In Kerkut, G. A. & 
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Received for publication 5 November 1994; revised and accepted 8 March 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 83-90 


NOTES ON THE MALE GENITALIA OF THE ANAEA RYPHEA - 
ANAEA EURYPYLE COMPLEX (NYMPHALIDAE) 


ASTRID CALDAS 


Dept. Biologia Animal e Vegetal - IB, Universidade do Estado do Rio de Janeiro, 
20559-900 Rio de Janeiro, R.J., Brazil 


ABSTRACT. Anaea ryphea resembles closely Anaea eurypyle, and both are found 
over the same geographic range. Separation of the two species has been based on two ex- 
ternal characters that vary continuously and unimodally. Genitalic dissections of 20 males 
with the traditional A. ryphea wing pattern and 20 males with the A. eurypyle wing pat- 
tern showed that male genitalic characters vary similarly in both taxa. There appears to be 
no consistent association between male genitalia and wing pattern variation in the A. 
ryphea - A. eurypyle complex. I conclude that the genitalic characters within this com- 
plex vary greatly and that no consistent “ensemble” exists that separate the taxa called A. 
ryphea and A. eurypyle, and these two “species” seem to be nothing but artificially desig- 
nated variants along gradients of continuous variation within a single, geographically 
widespread, species. 


Additional key words: Fountainea, intraspecific variation, Memphis, wing pattern. 


The genus Anaea (sensu lato) is very confusing and confused (see 
D’Abrera 1988). There is no cladistic treatment of it, and several of its 
species need careful reexamination. In the comprehensive revision of 
Comstock (1961) the genus contained 119 species, distributed in several 
subgenera. The species that were then assigned to subgenus Memphis 
are currently in three genera: Anaea, Memphis (DeVries 1987), and 
Fountainea (Rydon 1971, D’Abrera 1988). 

Anaea ryphea Cramer (=Memphis ryphea, =Fountainea ryphea) re- 
sembles closely Anaea eurypyle C. and R. Felder (=Memphis eurypyle, 
=Fountainea eurypyle) (Caldas 1994). They occur over similar geo- 
graphic ranges, from Mexico to Argentina and southern Brazil, although 
according to Comstock (1961) the two taxa overlap only from Mexico to 
Bolivia. He had no records of A. eurypyle from the Amazonian region 
or Brazil, but specimens from these regions can be found in other col- 
lections (A. Caldas, pers. obs.). Although the two species have been sep- 
arated by external characters, analyses of 499 males from localities 
throughout their geographic range showed that the two main external 
characters used to distinguish the species (the length of the “tail” on the 
hind wing and the degree of irregularity of the “mesial” line on the un- 
derside of the hind wing) vary in a continuous and correlated way, but 
with unimodal frequency distributions (Caldas 1996). One extreme of 
these distributions—long tail and straight “mesial” line—diagnoses the 
species A. eurypyle, and the other extreme plus the mode—short or no 
tail, irregular “mesial” line—diagnoses A. ryphea. However, many in- 
termediate states exist. This variation is suggestive of a single species. 

According to Comstock (1961), the male genitalic armature is consis- 


84 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Original drawings of (a) Anaea ryphea and (b) Anaea eurypyle male genitalia, 
after Comstock (1961). Scale bar = 1 mm. 


tently different between the two species. He provided line drawings of 
their genitalia (Fig. 1) to illustrate the main differences in the shape of 
the gnathos, valvae, aedeagus, and processes of the tegumen, but gave 
no further details in the text. Previously, however, Johnson and Com- 
stock (1941) had stated that “the structure of the gnathos in ryphea sep- 
arates it from all others of the group. The presence of tubercules [sic] in 


the central surface is unique.” 
Since the external morphological characters previously examined by 


VOLUME 51, NUMBER Il 85 


me (Caldas 1996) could not be used for distinguishing taxa in this com- 
plex, because of their unimodal distribution, I sought to determine 
whether genitalic characters could distinguish species. I compare my 
findings with the drawings in Comstock (1961). 


MATERIALS AND METHODS 


I dissected the genitalia of 20 male specimens with the wing pattern 
characteristic of A. ryphea and 20 specimens with the A. eurypyle pat- 
tern from the collection of the National Museum of Natural History, 
Smithsonian Institution. Specimens were from Peru, Brazil, Bolivia (both 
taxa), Colombia, Panama (A. ryphea), Mexico, Honduras, and Costa Rica 
(A. eurypyle). There were no individuals representative of the whole geo- 
graphic range for either species. My goal was to identify which struc- 
tures, if any, could distinguish the two taxa. Dissections were made in 
water, under a stereomicroscope, after washing the separated abdomens 
in alcohol (EtOH) and boiling them for 3.5 minutes in 10% potassium 
hydroxide (KOH). Genitalia were kept in vials with glycerine. 


RESULTS AND DISCUSSION 


Male genitalic characters vary similarly in both taxa, and no consistent 
trend was observed for the structures that Comstock (1961) used to 
separate A. ryphea and A. eurypyle. No two individuals with identical 
genitalia were found among the 40 males dissected. Some of the varia- 
tion is illustrated in Figs. 2 and 3, which show randomly selected geni- 
talia. These are drawings made in the same schematic way of the origi- 
nal drawings of Comstock (1961), in order to facilitate comparison with 
Fig. 1. Comparing the genitalia of three individuals with A. ryphea ex- 
ternal characteristics (Figs. 2a, 2b, and 2c) with Comstock’s drawing 
(Fig. la), the latter appears to be inaccurate. No individual with an A. 
ryphea wing pattern was found to have a small ventral spine on the 
tegumen, anterior to the gnathos; all had it long, as in Fig. lb. The 
gnathos did not present the shape illustrated in Fig. la, nor did the val- 
vae. Similarly, the aedeagus and saccus varied in shape and size 
throughout the complex (Figs. 2a, b, and c, no two aedeagi or sacci with 
the same shape). 

The genitalia in Figs. 3a, 3b and 3c cannot be considered different 
from those in Fig. 2, although they all belong to individuals with the A. 
eurypyle wing pattern. Again, they do not agree with Comstock’s drawing 
of A. eurypyle genitalic armature (Fig. 1b). No individual has the slender 
gnathos, the valvae vary in shape and length, as does the aedeagus (Figs. 
3a, b, and c). They bear the same long spine-like process of the tegumen 
shown in Figs. 2a, b, and c. In fact, the genitalia in Figs. 2 and 3 seem 
to be a mixture of characteristics from both Comstock’s drawings. 


86 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Poe pee 


Fic. 2. Genitalia of male individuals with Anaea ryphea wing pattern, respectively 
from (a) Panama, (b) Peru, (c) Brazil. Scale bar = 1 mm. 


Part of the difference seen by Comstock in the genitalia of A. ryphea 
and A. eurypyle may be due to the angle from which the genitalia were 
seen. He probably used slides of genitalia (F. Rindge, pers. comm.) to 
make his drawings, and slide mounting is likely to alter the shape of 
genitalia. Figs. 4 and 5 show photographs of the same genitalia from 


VOLUME 51, NUMBER 1 87 


a. - 
see 
meg 
ae 
i 
? 


a 
Fic. 3. Genitalia of male individuals with Anaea eurypyle wing pattern, respectively 
from (a) Bolivia, (b) Brazil, (c) Mexico. Scale bar = 1 mm. 


Figs. 2 and 3, taken from an angle different from the one used for the 
drawings (all drawings were made with the genitalia lying flat so that 
the superior or left side matched the inferior or right side). Thus, the 
gnathos appears slender (4c and 5a) or broad (4a and 5c). The uncus can 
appear shorter (5a), the same length (4a) or longer than the tegumen 
(5b). The tegumen itself always bears a long spine-like process beneath 
the gnathos, although Comstock’s drawing for A. ryphea (Fig. 1a) shows 


88 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 4-5. Male genitalia photographs. 4, male genitalia from individuals with Anaea 
ryphea wing pattern; 4a, b, and c (top to bottom) refer to drawings 2a, b, and c respec- 
tively. 5, male genitalia from individuals with Anaea eurypyle wing pattern; photographs 
5a, b, and c (top to bottom) refer to drawings 3a, b, and c respectively. Scale bar = 1 mm. 


VOLUME 51, NUMBER 1 89 


Fic. 6. Genitalic armature 3c/5c shown from five different angles (6a to 6e, starting 
on top left). Scale bar = 1 mm. 


a small process. Shape and size of the valvae vary greatly. Again, the six 
genitalia in Figs. 4 and 5 show that no two valvae are completely similar. 

While taking the previous photographs, I noticed that a slightly dif- 
ferent angle sometimes provides very different views of the same 
armature. To further illustrate my point, I decided to have photos of the 
same genitalic armature taken from different angles. Thus, Figs. 6a—6e 


90 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


are all from the same armature shown in Figs. 3c and 5c. As the angle 
varies, the gnathos can appear more (6a, 6b, 6e) or less slender (6c, 6d), 
the proportion of the uncus in relation to the tegumen varies—tegumen 
longer than the uncus, Fig. 6c (compare with 1b), or shorter than the 
uncus, Figs. 6b, 6d (compare with Fig. la)—and the aedeagus can ap- 
pear as in Fig. la (Fig. 6c) or 1b (Fig. 6d), both Comstock’s illustrations. 
This reinforces my idea that the variation shown in the latter may be 
partially due to the angle from which the genitalia were seen. Of course, 
Figs. 6a and 6e are extremes, but they show very well how slender the 
gnathos can appear. 

I conclude that the genitalic characters within this complex vary 
greatly and that no consistent “ensemble” exists that separate the taxa 
called A. ryphea and A. eurypyle. These results, together with the re- 
sults of my previous studies (Caldas 1996), suggest that these two “spe- 
cies” are nothing but artificially designated variants along gradients of 
continuous variation within a single, geographically widespread, spe- 
cies. Another species in the group—Anaea ecuadoralis, which resem- 
bles A. ryphea and A. eurypyle closely in many features—may also be 
part of this variable species. 


ACKNOWLEDGMENTS 


I thank the Smithsonian Institution, which, through its curators, allowed the develop- 
ment of this research. John M. Burns, Donald Harvey and Robert K. Robbins offered sug- 
gestions and held helpful discussions throughout the project. I especially thank Elizabeth 
Klafter, who provided most logistic and technical support I needed, and Carl C. Hansen, 
for that extra effort with the photographs. This research was funded by two CAPES/Min- 
istry of Education/Brazil grants, a Smithsonian short-visit grant, and several SR-2/Univer- 
sidade do Estado do Rio de Janeiro fundings. 


LITERATURE CITED 


CaLpas, A. 1994. Biology of Anaea ryphea (Nymphalidae) at Campinas, Brazil. J. Lepid. 
Soc. 48:248—257. 

. 1996. Intraspecific variation of Anaea ryphea and Anaea eurypyle (Nymphali- 
dae). J. Res. Lepid. 32:37—44. 

Comstock, W. P. 1961. Butterflies of the American tropics. The genus Anaea, Lepi- 
doptera, Nymphalidae. American Museum of Natural History, New York. 214 pp. 
D’ABRERA, B. 1988. Butterflies of the neotropical region. Part V. Nymphalidae (Conc.) & 

Satyridae. Hill House Publ. 

DEVRIES, P. J. 1987. Butterflies of Costa Rica and their natural history. Princeton Univer- 
sity, New Jersey. 

JoHNSON, F. & W. P. Comstock. 1941. Anaea of the Antilles and their continental rela- 
tionships with descriptions of new species, subspecies and forms (Lepidoptera, 
Rhopalocera, Nymphalidae). J. New York Entomol. Soc. 49:301—343. 

MUYSHONDT, A. 1974. Notes on the life cycle and natural history of butterflies of El Sal- 
vador. VI. Anaea (Memphis) eurypyle confusa (Nymphalidae). J. Lepid. Soe. 
28:306-—314. 

RypDon, A. H. B. 1971. The systematics of the Charaxidae (Lepidoptera: Nymphalidae). 
Entomol. Rec. J. Var. 83:339—341. 


Received for publication 2 September 1995; revised and accepted 19 February 1996. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
51(1), 1997, 91-92 


BODY WEIGHT AS RELATED TO WING MEASURE IN 
HAWKMOTHS (SPHINGIDAE) 


Forewing or hindwing length (L) and forewing span (S) are often used as species body- 
size indices in Lepidoptera, but the body weights (W) actually represented are not usually 
known. Establishing mathematical relations between total body weight or mass and wing 
measure among species of a taxon not only validates the use of wing measure as a body 
size surrogate (Miller 1996), but also enables estimation of body weight from wing mea- 
sure for physiological and ecological purposes. In detail, such relations are specific for dif- 
ferent groups; in form, they are usually allometric power functions of W = a (L> or S>) or 
the logarithmic equivalent (Miller 1977). Wing length is defined here as the distance be- 
tween the wing base (excluding tegula) and the farthest extending wing tip (including 
fringe); wing span is defined as the farthest distance between the wing tips on specimens 
spread in the usual manner. 

In their studies of flight mechanics, Bartholomew and Casey (1978) and Casey (1976) as- 
sembled and tabulated W and L values for 15 and 38 identified hawkmoth species, respec- 
tively. Although they did not derive relations between W and L in the foregoing papers, 
Casey (1989) presented such a relation later. This relation had L rather than W as the de- 
pendent variable, was based on only the smaller of the above data sets, and was displayed on 


Live body wt. 
(g) (W) 


W = 0.000548 (L*°°) 
Raw r2=0.90 


© Macroglossinae 
e Sphinginae 


0 20 40 60 80 100 
Forewing length (mm) (L) 
Fic. 1. Live body weight as related to forewing length in hawkmoths. Each point rep- 


resents from one to several individuals of either or both sexes of a species. Data from 
Bartholomew and Casey (1978) and Casey (1976). 


92 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


a double-logarithmic scale. Here I recharacterize the relation with W as the dependent vari- 
able using both of the above data sets and displaying results on an arithmetic scale. 

Study data here are 70 observations on live weight (W), forewing length (L), and 
forewing span (S) of field-caught individuals of 48 identified species (Bartholomew & 
Casey 1978, Casey 1976). These species include 33 Macroglossinae and 15 Sphinginae— 
the sole subfamilies recognized in Sphingidae (Hodges 1971, Pittaway 1993)—and there 
are 29 New World and 19 Old World species. The sexes are pooled because many individ- 
uals originally were not sexed. Species live weights range from 0.18 to 6.25 g, essentially 
the full range for the family. For display, two or more observations for the same species 
were averaged. In analysis, species means were weighted by the number of observations. 
Where only S was originally given for a species, as in the data set of Casey (1976), I de- 
rived L after measuring L and S on D’Abrera’s (1986) life-size illustrations such that L = 
S(Lp:abrera/SprAbrera): LO solve the equation W = a (L>), I used the quasi-Newton loss func- 
tion for ordinary least-squares nonlinear regression (SYSTAT 1992). I derived proportion- 
ality functions for interconverting L and S by simple ordinary least-squares regression. 

The recharacterized family-level relation between live body weight (W) and forewing 
length (L) is W = 0.000548 (L?2.%) (Fig. 1). It is evident from Fig. 1 that sample species of 
Sphinginae are larger in mean size than those of Macroglossinae, an outcome that may re- 
flect a real condition among hawkmoths. The subfamilies do not seem to differ in the W 
versus L relation, however. The proportionality functions which serve to estimate L from 
S and vice versa when one or the other measurement is available but the other desired are 
L = -1.09 + 0.467 S, and S = 4.46 + 2.094 L (r? = 0.98). 

This study reconfirms that hawkmoth body weight is indexed by wing measure. Live body 
weight increases as the square of forewing length in Sphingidae, whereas it increases as the 
cube in Tortricidae (Miller 1977). The difference is fundamental, and reflects the empirical 
nature of such relations and their dependence on morphometrics and physiology. 

Live weights estimated from the nonlinear equation with forewing length (Fig. 1) are 
not intended for rigorous use. Rigorous use would require not only confidence intervals 
for the equation, but also body-weight controls in the original sources for sex, egg load, 
and crop content. 

I thank L. S. Fink, T. M. Casey and an anonymous reader for useful review comments 
on the manuscript. 

LITERATURE CITED 


BARTHOLOMEW, G. A. & T. M. CASEY. 1978. Oxygen consumption of moths during rest, 
pre-flight warm-up, and flight in relation to body size and wing morphology. J. Exp. 
Biol. 76:11—25. 

CasEy, T. M. 1976. Flight energetics of sphinx moths: power input during hovering flight. 
J. Exp. Biol. 64:529-543. 

. 1989. Oxygen consumption during flight, pp. 257-272. In Goldsworthy, G. J. & 
C. H. Wheeler (eds.), Insect flight. CRC Press, Boca Raton, Florida. 

D’ABRERA, B. 1986. Sphingidae mundi: hawk moths of the world. E. W. Classey, Faring- 
don, United Kingdom. 226 pp. 

HopcEs, R. W. 1971. Sphingoidea: hawkmoths. Moths of America North of Mexico Fasc. 
21. E. W. Classey & R. B. D. Publications, London. 158 pp. 

MILLER, W. E. 1977. Wing measure as a size index in Lepidoptera: the family Olethreu- 
tidae. Ann. Entomol. Soc. Am. 70:253—256. 

. 1997. Diversity and evolution of tongue length in hawkmoths (Sphingidae). J. 
Lepid. Soc. 51:9-31. 

Pirraway, A. R. 1993. The hawkmoths of the western Palaearctic. Harley Books, Col- 
chester, Essex, England. 240 pp. 

SYSTAT. 1992. Statistics, version 5.2 edition. SYSTAT Inc., Evanston, Illinois. 724 pp. 


WILLIAM E. MILLER, Entomology Department, University of Minnesota, Saint Paul, 
Minnesota 55108, USA. 


Received for publication 10 May 1995; revised and accepted 6 January 1996. 


VOLUME 51, NUMBER 1 93 


Journal of the Lepidopterists’ Society 
51(1), 1997, 93-95 


OCCURRENCE OF THE PALAEARCTIC MOTH, CNEPHASIA LONGANA 
(TORTRICIDAE), ON SANTA ROSA ISLAND, CALIFORNIA 


Additional key words: omnivorous leaf-tier, larval competition, Asteraceae, Lami- 
aceae, Scrophulariaceae. 


Santa Rosa Island is the second largest of the eight California Channel Islands, but its 
Lepidoptera fauna has been poorly known relative to most of the other islands, primarily 
due to its long term private ownership. Only about 125 species of Lepidoptera were 
recorded, perhaps 30—40% of an expected total (Powell 1994). Hence, I was not surprised 
to find 120+ additional species during my first visit, in late April to early May 1995. One 
of the previously unrecorded species I did not expect was Cnephasia longana (Haworth) 
(Tortricidae: Tortricinae), a Palaearctic species that has been known in coastal central Cal- 
ifornia since the 1940s (Powell 1964) but not on any of the Channel Islands. 

Cnephasia longana, the so-called ‘omnivorous leaf-tier, was reported in North America 
in the Pacific Northwest beginning in 1929, where it became a widespread pest of field 
crops. It was first recognized in California in 1947 (Keifer 1948); within a few years C. lon- 
gana was found widely in the San Francisco and Monterey Bay areas, associated with cul- 
tivated and native flowers, strawberries, and other field crops (CDFA records, Mid- 
dlekauff 1949, Powell 1964). 

The species is univoltine; winter is passed by first instar larvae in silken hibernacula, 
and populations are most easily detected by late instar larvae in April and May or adults, 
which are attracted to lights, from late April to early July, varying with year and locality. 
First and second instar larvae are leaf miners in low herbs; later instars web terminal parts 
of the plants, particularly buds and flowers. The species is polyphagous and in California 
most often feeds on Asteraceae and other herbaceous plants such as poppies, flax, and cul- 
tivated flowers, in open field situations (Powell 1964). 

Subsequently, C. longana has extended its range in California, having been discovered 
in the Humboldt Bay area (1960), Lake County (1969), San Benito County (1964), and 
southward, in coastal San Luis Obispo County (1967) and Santa Barbara County at Lompoc 
(1970) and Goleta (1976) (CDFA records, Essig Museum, R. Priestaf specimens) (Fig. 1). 

Scott Miller collected Lepidoptera, including some tortricoids, on Santa Rosa Island in 
April 1976 and May 1978 and did not take C. longana (LACM and SBNHM records). 
Negative evidence suggests this adventive species was not established elsewhere on the 
Channel Islands then and in the early to mid 1980s. We did not find adults or larvae on 
several of the islands: on Santa Cruz Island in May 1984, when a group of four lepidopter- 
ists surveyed for microlepidoptera during a four-day visit; C. Drost made many moth col- 
lections on Anacapa and Santa Barbara islands during 1985-1988; on the southern islands, 
we sampled on San Nicolas and Santa Catalina in May 1978; S. Bennett collected exten- 
sively on Catalina in 1981-1982; and I collected for a week on San Clemente in April 1980. 

On Santa Rosa Island I found males of C. longana flying at dawn (0600 PST at 8°C) on 
the first morning of my visit, and adults appeared abundantly during the following week. 
They occurred from sea level to the highest peaks (460—480 m), in cattle grazed fields 
from the east coast to the deflation plane back of the most western coastal dunes. Adults 
were taken in most blacklight samples, up to 85 in a single trap. 

Competitive exclusion by introduced species is a subject that has not been investigated 
in detail by lepidopterists but may play a role in decline of native species that are near 
ecological homologues of sympatric immigrants. On Santa Rosa Island I found larvae of 
C. longana numerous in flower heads of four species of Asteraceae: the weed Achillea 
millefolium; a native Cirsium, competing with Platyptilia carduidactyla (Riley) (Ptero- 
phoridae), which probably is introduced on the islands; Erigeron glauca, competing with 
Platyptilia williamsi Grinnell; and in two species of Gnaphalium, competing with Tebenna 
gnaphaliella (Kearfott) (Choreutidae), Phaneta apacheana (Wlsm.) (Tortricidae), Pata- 
gonia peregrina (Heinrich) (Pyralidae), and Platyptilia williamsi. The last three moth 


94 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


wan ae ane === 


, 
£ etter 


Pe Miles 


200 Kilometers 


Fic. 1. Distribution of Cnephasia longana in California: cross-hatched area indicates 
the known range in the San Francisco and Monterey Bay areas by 1962; dated localities 
(solid dots) refer to earliest known records in other areas, and Santa Rosa Island (stip- 


pled). 


species were encountered commonly on Santa Rosa, while T: gnaphaliella was very rare (2 
adults), whereas it was abundant on San Clemente Island in April 1980. Larvae of C. lon- 
gana also fed in flowers of Castilleja affinis (Scrophulariaceae) , where they are potential 
competitors with Schreckensteinia felicella (Wlsm.) (Schreckensteiniidae), Endothenia 
hebesana (Wlk.) (Tortricidae), and Amblyptilia pica (Wlsm.) (Pterophoridae), which are 
inhabitants of this plant in coastal central California but are not known on the island. I also 
found larvae of C. longana on Stachys bullata (Lamiaceae), the host of Endothenia condi- 
tana in the San Francisco Bay area. 

The omnivorous leaf-tier is a potential competitor with two endemic insular Lepi- 
doptera, Argyrotaenia franciscana insulana Powell (Tortricidae) and Euphydryas editha 


VOLUME 51, NUMBER 1 95 


insularis Emmel & Emmel (Nymphalidae). The former is polyphagous, and I collected its 
larvae on Achillea and Erigeron on Santa Rosa. The butterfly is a specialist on Scrophular- 
iaceae, and I found the young larvae on Castillela exserta |=Orthocarpus purpurascens], 
which is a likely host of C. longana. 

I thank C. Mack Shaver, Superintendent, for issuing a permit to inventory Lepidoptera 
in the Channel Islands National Park, and personnel of the Resource Division, whose co- 
operation facilitated my visit: Tim Coonan, Heide David, CeCe Sellgren, and David 
Kushner, particularly the last, who assisted with collections. Tom Eichlin provided copies 
of early records in the Calif. Dept. Food & Agric., Plant Pest Diagnostics Branch, Sacra- 
mento (CDFA records); cooperation by curators of the Los Angeles County Museum 
(LACM) and Santa Barbara Natural History Museum (SBNHM) enabled use of speci- 
mens; and Richard Priestaf provided records of Santa Barbara microlepidoptera. 


LITERATURE CITED 


KEIFER, H. 1948. Systematic entomology, pp. 205—209. In Armitage, Ann. Rept. Bureau 
Entomol. & Plant Quarantine, Calif. Dept. Agric. Bull. 37. 

MIDDLEKAUFF. 1949. The omnivorous leaf tier in California. J. Econ. Entomol. 42:35-36. 

POWELL, J. A. 1964. Biological and taxonomic studies on tortricine moths, with reference 
to the species in California. Univ. Calif. Publ. Entomol. 32:1—317. 

. 1994. Biogeography of Lepidoptera on the California Channel Islands, pp. 

449-464. In Halvorson, W. & G. Maender (eds.), The fourth California Island sym- 

posium: update on the status of resources. Santa Barbara Mus. Nat. Hist., Santa Bar- 

bara, California. 


J. A. POWELL, Essig Museum of Entomology, University of California, Berkeley, Califor- 
nia 94720, USA. 


Received for publication 20 October 1995; revised and accepted 12 March 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 95-97 


EFFECTS OF GENE-ENVIRONMENT INTERACTION ON SILK YIELD IN 
ANTHERAEA MYLITTA (SATURNIIDAE) 


Additional key words: tasar silk moth, absolute silk yield, Terminalia arjuna, stability. 


Antheraea mylitta (Drury) is a saturniid moth of considerable commercial value used 
for production of tasar silk. Because interactions between genotype and environment may 
exert significant influence over specific life history features (Falconer 1952, Dickerson 
1962, Hanson 1964, Breese 1969), it is likely that silk yield and yield contributing traits in 
different strains of A. mylitta are influenced by seasonal and/or environmental factors 
(Jolly et al. 1979). In an effort to understand features that may contribute to the maxi- 
mization of silk production, we conducted rearing experiments to measure the interaction 
between genotype and environment for silk yield and to screen stable genotypes of A. 
mylitta for use in breeding programs to enhance silk yield. 

We investigated eight diverse genetic strains of A. mylitta: Nagri,, Nagri,, Nagris, 
Sukli, Raily, Sukinda, Laris (P), and Palma. The genotype lines were obtained from the 
germplasm bank of the Central Tasar Research and Training Institute, Ranchi, Bahir, In- 
dia. We reared the eight genotypes through two generations in July-August and Octo- 
ber—November of 1988. The two generations mature under different environmental con- 
ditions: the July-August brood occuring during the rainy season, and the October— 
November brood occuring during the dry season. Larvae were reared on individual plants 


96 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Analysis of variance (ANOVA) for silk yield and related parameters in Bom- 
byx mori as a function of genotype and environment. All parameters in the table are sig- 
nificant at P < 0.01. 


Mean sums of squares 


Absolute Larval Larval Cocoon Shell Shell 
Source silk yield Fecundity weight span E.R.R.% weight weight ratio 
G (genotype) 9586.55 12288.29 28.20 9.30 248.93 17.24 0.28 3.16 


E (environment) 83665.97 48260.00 353.27 105.02 737.35 174.57 22.62 533.27 
G x E interaction 5663.00 5637.23 21.61 7.09 615.21 2.89 014 3.50 


of Terminalia arjuna Bedd. (Combretaceae) situated in rows, with each plant separated 
by at least 2 m. Experimental design of the rearings followed a randomized block strategy 
during both generations, with three replicates of each genotype, 300 larvae per replica- 
tion. Absolute silk yield was estimated based on shell weight of all the cocoons harvested 
from each replication. Gene-environment interaction was calculated following the 
methodology proposed by Plaisted and Peterson (1959). Analysis of variance (ANOVA) 
was calculated by pooling absolute silk yield of the two seasons. 

Table 1 indicates that there were significant differences among genotypes (G), environ- 
ments (E), and in the gene-environment interaction, suggesting that genotypes interact 
considerably with environmental conditions to produce different silk yields. Mean ab- 
solute silk yield for the two generations and within generation type variance (62VL or sta- 
bility) are illustrated in Table 2. Mean absolute silk yield ranged from 25.07 g to 147.78 g 
in the first generation, while that of the second generation ranged from 67.08 g to 259.73 
g, illustrating a marked between generation difference. Absolute silk yield was found to 
be much higher in all genotypes during the second generation, corroborating the findings 
of Jolly et al. (1979). 

As illustrated in Table 2, the genotypes in order of increasing absolute silk yield in the 
first generation were Palma (25.07 g), Laria (49.49 g), Sukinda (61.46 g), Raily (82.65 g), 
Nagri, (96.70 g), Sukli (98.98 g), Nagri, (100.64 g), and Nagri, (147.78 g). In the second 
generation, absolute silk yield (from least to greatest) was achieved by Laria, Nagris, 
Palma, Sukinda, Raily, Sukli, Nagri,, and Nagri,. 

The estimate of genotype x generation variance exhibited a wide range from 792.96 to 
4280.26 (Table 2). The greatest between-generation variability was demonstrated by Na- 
gri, followed by Palma, Nagri,, and Nagri,. The least between-generation variability was 
demonstrated by Raily, followed by Sukli and Sukinda. These results suggest that greater 
stability in silk yield (between generations) could be obtained from Raily. Hence, this 
genotype would respond better to between-generation differences because of the lesser 
influence of environment on its absolute silk yield. 


TABLE 2. Mean absolute silk yield and stability for eight genotypes of Bombyx mori 
reared in different environments. 


Absolute silk yield in environment 
GxE 
Genotypes Rainy season Dry season interaction 
Nagri, 96.70 259.73 2704.48 
Nagri, 100.64 234.91 2819.07 
Nagri, 147.78 123.63 4280.26 
Sukli 96.98 181.01 ie SOSA 
Raily 82.65 157.58 792.96 
Sukinda 61.46 156.66 991.73 
Laria (P) 49.49 67.08 2190.97 


Palma 25.07 148.01 3585.83 


VOLUME 51, NUMBER 1 97 


LITERATURE CITED 


BREESE, E. L. 1969. The measurement and significance of genotype environment interac- 
tions in grasses. Heredity 24:27—44. 

DICKERSON, G. E. 1962. Implications of genetic environmental interaction in animal 
breeding. Animal Pred. 4:47—64. 

FALCONER, D. S. 1952. The problem of environment and selection. Am. Nat. 86:293—298. 

HANSON, W. D. 1964. Genotype environment interaction concepts for field experiments. 
Biometrics 20:540—552. 

JOLLy, M. S., S. K. SEN, T. N. SONWALKAR & G. K. PRASAD. 1979. Non-mulberry_ silks. 
Food and Agriculture Organization of the United Nations, Rome. 178 pp. 

PLAISTED, P. L. & L. C. PETERSON. 1959. A technique for evaluating the ability of selec- 
tion to yield consistently in different locations or seasons. Am. Potato J. 36:381—385. 


A. K. SENGuPTA, Silkworm Breeding and Genetics Section, Central Sericultural Re- 
search and Training Institute, Berhampore 742-101, West Bengal, India, AND A. A. SID- 
DIQUI, Regional Muga Research Station, Boko, 781-123, Assam, India. 


Received for publication 4 January 1993; revised and accepted 13 May 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 97-101 


DISTRIBUTION OF A NORTHERN FAUNA OF NOCTUIDAE IN THE 
MOUNTAINS OF OREGON 


Additional key words: endemism, non-target species, biogeography. 


Although the Oregon butterfly fauna has been well studied (Dornfeld 1980), compara- 
tively little was known about the Oregon moth fauna until about 1960. During the past 30 
years, extensive collecting has been conducted in the state, most notably by Stanley G. 
Jewett, Jr, C. William Nelson, James H. Baker, Elmer L. Griepentrog, Victor B. 
McHenry, Kenneth J. Goeden, and Jeffrey C. Miller. From 1992 through 1995, the U. S. 
Forest Service also conducted extensive blacklight (UV) trap sampling in the Cascade 
Range and the Blue Mountains to assess the impacts on nontarget Lepidoptera of Bacillus 
thuringiensis subsp. kurstaki sprays for suppression of outbreaks of western spruce bud- 
worm (Choristoneura occidentalis Freeman: Tortricidae) (see Grimble et al. 1992 for de- 
tails of the sampling protocols in these studies). 

Various components of this moth fauna show biogeographic connections with the 
northern Pacific Coast, California, the Great Basin, and the northern Rocky Mountains. 
In this paper, we report on a northern fauna of Noctuidae that is transcontinental across 
Canada from Quebec to British Columbia, extending southward through the Appalachi- 
ans to North Carolina, the Rocky Mountains, and the mountains of Oregon. The fauna 
has been enumerated by Rockburne and Lafontaine (1976), Prentice (1962) and from a 
survey of museum records. Only those species typical of northern hardwood-conifer 
forests, meadows, or wetlands are included in this study; ubiquitous and/or migratory spe- 
cies throughout most of North America, such as Heliothis zea (Boddie) and Peridroma 
saucia (Hbn.), are excluded from consideration. 

This northern noctuid fauna is largely confined to three mountainous regions of Ore- 
gon; the northern Coast Range, the Cascade Range, and the Blue Mountains. The north- 
ern Coast Range consists of low mountains 300-600 m in elevation, with a few higher 
peaks to 900 m, extending from Clatsop County to coastal Lane County. The Cascade 
Range extends from Multnomah and Wasco Counties south to Jackson and Klamath 
Counties. The lower Cascade foothills of the western slope extend from 150-1500 m, 
whereas the high Cascades are 1200-2100 m in elevation, with high volcanic peaks over 
3000 m. The Blue Mountains extend from Crook County northeast to Wallowa County, 


98 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Distribution of northern species of Noctuidae in the Coast Range (CO), 
Cascade Range (CA), and the Blue Mountains (BM) of Oregon. X = collected, O = not 
collected). 


(@) 
e) 
Q 
> 
w 
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Species 


Abrostola urentis Gn. 
Acronicta fragilis Gn. 
Acronicta funeralis G.& R. 
Acronicta grisea Wk. 
Acronicta impleta Wk. 
Acronicta impressa Wk. 
Acronicta innotata Gn. 
Acronicta lepusculina Gn. 
Acronicta radcliffei (Harv.) 
Agroperina inficita (WIk.) 
Agrotis obliqua (Sm.) 

Agrotis venerabilis Wk. 
Aletia oxygala (Grt.) 

Alypia langtoni Couper 
Amphipoea americana (Speyer) 
Amphipyra pyramidoides Gn. 
Amphipyra tragopoginis (Cl.) 
Anaplectoides prasina D.& S. 
Anaplectoides pressus (Grt.) 
Anathix puta (G. & R.) 
Androloma maccullochi (Kby.) 
Anhimella contrahens (Wlk.) 
Apamea alia (Gn.) 

Apamea amputatrix (Fitch) 
Apamea finitima Gn. 
Apamea impulsa (Gn.) 
Apamea indocilis (WIk.) 
Apamea inordinata (Morr.) 
Apamea lignicolora (Gn.) 
Apamea plutonia (Grt.) 
Apamea vultuosa (Grt.) 
Aplectoides condita (Gn.) 
Autographa ampla (Wlk.) 
Autographa mappa (G.& R.) 
Autographa pseudogamma (Grt.) 
Autographa sansoni Dod 
Brachylomia algens (Grt.) 
Catabena lineolata Wlk. 
Catocala briseis Edw. 
Catocala relicta Wlk. 
Catocala semirelicta Grt. 
Chersotis juncta (Grt.) 
Chortodes inquinata (Gn.) 
Chortodes rufostrigata (Pack.) 
Cosmia calami (Harv.) 
Cucullia florea Gn. 

Cucullia intermedia Speyer 
Cucullia omissa Dod 
Cucullia postera Gn. 
Cucullia speyeri Lint. 
Diachrysia aeroides (Grt.) 


XOOKOKXOOOOKK MOK KOK KOK K KOK KM MK KMOKOOKK KKM AKXOOCOKOOK KK KK MK 
~ MK OM mK KO KOK KK KK KOKO KOM MK KKM MK MMM KKK MOKOOKKK AKAM 
KKK MH MOKOKM KM KOK OKOK MK KMOOKM KOM MK MMMM MAMOOCOKMKKKOKMKAKMKAXORKO 


VOLUME 51, NUMBER 1 


Species 


Diarsia rosaria (Grt.) 
Egira dolosa (Grt.) 
Enargia decolor (Wlk.) 
Enargia infumata (Grt.) 


Eosphoropteryx thyatyroides (Gn.) 


Eremobina claudens (WIlk.) 
Eueretagrotis perattenta (Grt.) 
Euplexia benesimilis McD. 
Eupsilia tristigmata (Grt.) 
Eurois astricta Morr. 

Eurois occulta (L.) 
Eutricopis nexilis (Morr.) 
Euxoa declarata (Wlk.) 
Euxoa divergens (WIk.) 
Feralia comstocki (Grt.) 
Galgula partita Gn. 
Graphiphora haruspica (Grt.) 
Hecatera sutrina (Grt.) 
Heliothis phloxiphaga G.& R. 
Helotropha reniformis (Grt.) 
Homorthodes furfurata (Grt.) 
Hyppa xylinoides (Gn.) 
Lacanobia grandis (Gn.) 
Lacanobia lilacina (Harv.) 
Lacanobia lutra (Gn.) 
Lacanobia nevadae (Grt.) 
Lacanobia radix (Wlk.) - 
Lacanobia subjuncta (G.& R.) 
Lacanobia tacoma (Stkr.) 
Lacinipolia olivacea (Morr.) 
Lacinipolia vicina (Grt.) 
Leucania insueta Gn. 
Lithacodia albidula (Gn.) 
Litholomia napaea (Morr.) 
Lithomoia solidaginis (Hbn.) 
Lithophane amanda (Sm.) 
Lithophane baileyi Grt. 
Lithophane georgii Grt. 
Lithophane innominata (Sm.) 
Lithophane petulca Grt. 
Lithophane thaxteri Grt. 
Luperina passer (Gn.) 
Marathyssa inficita (WIk.) 
Melanchra adjuncta (Gn.) 
Melanchra pulverulenta (Sm.) 
Mniotype ducta (Grt.) 
Mniotype tenera (Sm.) 
Nephelodes minians Gn. 
Nycteola cinereana N.& D. 
Nycteola frigidana (WIk.) 
Ochropleura plecta (L.) 
Oligia illocata (Wlk.) 
Oncocnemis piffardi (Wlk.) 


TABLE 1. 


Q 
eo) 


OK KKOOKOKKOKOKK HOM KM MK KOK KKM HKMOKMKOKOKKOKMK KK KOOOKKXOOKOOORK 


@) 
> 


KK MM OO KM MM OM MO OO SO OO OO OS OS OS OO SOS OS OO OOO OS OO OM OO MS OS OM 


ey 
x 


OKOOK KOKO KK MMMM MOK KK OM KM MOM KK KKM OK MO KOKO KKK MK MK OM KM 


100 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Continued. 


Q 
ie) 
@ 
> 
w 
= 


Species 


Oncocnemis riparia Morr. 
Orthosia hibisci (Gn.) 
Orthosia revicta (Morr.) 
Orthosia segregata (Sm.) 
Papestra cristifera (Wlk.) 
Papestra quadrata (Sm.) 
Paradiarsia littoralis (Pack.) 
Phlogophora periculosa Gn. 
Platyperigea meralis (Morr.) 
Platypolia anceps (Steph.) 
Polia nimbosa Gn. 

Polia purpurissata (Grt.) 
Protolampra rufipectus (Morr.) 
Proxenus miranda (Grt.) 
Pyrrhia exprimens (WIk.) 
Raphia frater Grt. 

Rhyacia quadrangula (Zelt.) 
Scoliopteryx libatrix (L.) 
Sideridis maryx (Gn.) 
Sideridis rosea (Harv.) 
Spaelotis clandestina (Harr.) 
Stretchia plusiaeformis Edw. 
Synedoida adumbrata (Behr) 
Syngrapha alias (Ottol.) 
Syngrapha epigaea (Grt.) 
Syngrapha rectangula (Kby.) 
Syngrapha viridisigma (Grt.) 
Ufeus satyricus Grt. 

Xanthia togata (Esper) 
Xestia collaris (G.& R.) 
Xestia smithii (Snell.) 

Xylena cineritia Grt. 

Xylena curvimacula (Morr.) 
Xylena nupera (Lint.) 
Xylena thoracica (Putnam-Cramer) 
Xylotype acadia B. & Benj. 
Zale duplicata (Bethune) 
Zale lunata (Drury) 

Zale minerea (Gn.) 


xXKXOKOKOOKOKOOKKKMOOOKMOKOKKKKKMKMOOKOOKORKKO 
KX OK KM KK KK OK KKM OM KM MK MM KK OKMOOKOK KM 
OK KK KK KM MOK KOK KK KM MMMM MOK KOO KO KK MM HK OM 


and include the Ochoco, Strawberry, Elkhorn, Wallowa and Imnaha Mountain ranges. EL 
evations are circa 1200-1800 m, with high peaks of 2700-3000 m. Both the Coast Range 
and the Cascade Range generally have a cool, moist climate due to the influence of the 
Pacific Ocean, although it is drier in the rain shadows on the eastern slopes. The Blue 
Mountains have a dry, more severe continental climate similar to the northern Rocky 
Mountains of Idaho and Montana. 

To date a total of 143 of the 200 species of northern N octuidae with a transcontinental 
distribution across Canada have been collected in the mountains of Oregon. Table | out- 
lines the known distributions of these species in the Coast Range, Cascade Range, and 
the Blue Mountains. At present, we know of 707 species of Noctuidae recorded through- 


VOLUME 51, NUMBER 1 101 


TABLE 2. Number and percent of northern species of Noctuidae shared among the 
Coast Range (CO), Cascade Range (CA), and the Blue Mountains (BM) of Oregon. 


Species CoO CA BM CO/CA CA/BM CO/CA/BM 
Number 4 8 15 22, 34 60 


Percent 3 6 10 55 24 42, 


out the state, and so these northern taxa comprise about 20% of the total Oregon fauna. 
Table 2 indicates the number and percent of northern species shared among the three 
mountain regions. For species restricted to a single region, the Blue Mountains have the 
highest number (15) and the Coast Range the fewest (4). However, the Cascade Range 
has the highest total number of northern species with 124, followed by the Blue Moun- 
tains with 109 and the Coast Range with 86. 

This northern fauna is isolated and relictual in Oregon today, surviving on montane is- 
lands surrounded by broad expanses of desert lowlands and valleys. We postulate that 
these species dispersed southward into Oregon during the various glaciations of the Pleis- 
tocene. The modern distribution of endemically restricted species may suggest three dis- 
persal routes into the state during the Pleistocene. These include a northeastern route 
into the Blue Mountains from the adjacent mountains of northern Idaho and Montana, a 
north-south route through the Cascades from British Columbia, and possibly some dis- 
persal along the lower mountains of the Coast Range. Examples of such endemics include 
Orthosia segregata (Sm.), Papestra quadrata (Sm.), and Zale duplicata (Bethune) in the 
Blue Mountains; Lacanobia grandis (Gn.), Oncocnemis piffardi (Wlk.), and Lithophane 
baileyi Grt. in the Cascades; and Mniotype tenera (Sm.), Apamea impulsa (Gn.), and A. 
plutonia (Grt.) in the Coast Range. Moreover, it is significant that a number of northern 
species known from the Washington Cascades do not cross the Columbia River valley into 
the Oregon Cascades. These include Lithophane pexata Grt., L. fagina Morr., Anomog- 
yna speciosa (Hbn.), and A. perquiritata (Morr.). 

In conclusion, about 71% of the northern transcontinental noctuid fauna are known to 
occur in the mountains of Oregon. It is doubtful that many additional species of this fauna 
remain to be discovered in Oregon, with perhaps the exception of the high Wallowa 
Mountains where little survey work has yet been done. 

We thank Roy C. Beckwith, Lucille Clark, Roy Magelssen, and Dixie Moore for their 
help with field collecting and data. We are especially indebted to John D. Lattin and 
Richard Westcott for assistance with the insect collections at the Systematic Entomology 
Laboratory, Oregon State University, and the Oregon Department of Agriculture. Partial 
funding for this work was provided by the National Agricultural Pesticide Impact Assess- 
ment Program (NAPIAP), U. S. Forest Service. 


LITERATURE CITED 


DORNFELD, E. J. 1980. The butterflies of Oregon. Timber Press, Forest Grove, Oregon. 
276 pp. 

ernst 1. G., BECKWITH, R. C. & P. C. HAMMOND. 1992. A survey of the Lepidoptera 
fauna from the Blue Mountains of eastern Oregon. J. Res. Lepid. 31:83—102. 

PRENTICE, R. M. 1962. Forest Lepidoptera of Canada recorded by the forest insect sur- 
vey, vol. 2. Forest Entomol. Pathol. Branch, Canada Dept. Agric., Ottawa. 281 pp. 

ROCKBURNE, E. W. & J. D. LAFONTAINE. 1976. The cutworm moths of Ontario and Que- 
bec. Research Branch, Canada Dept. Agric., Ottawa. 164 pp. 


PAUL C. HAMMOND, Department of Entomology, Oregon State University, Corvallis, 
Oregon 97331, USA, AND DavID G. GRIMBLE, U.S.D.A Forest Service, Pacific Northwest 
Research Station, 3200 SW Jefferson Way, Corvallis, Oregon 97331, USA: 


Received for publication 10 January 1995; revised and accepted 1 April 1996. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 102 


W. D. WINTER COLLECTION TO THE MUSEUM OF COMPARATIVE 
ZOOLOGY, CAMBRIDGE 


Additional key words: Geometridae, Noctuidae, New England. 


In October 1996, the Lepidoptera collection of William D. (Dave) Winter (Westwood, 
Massachusetts) was donated to the Museum of Comparative Zoology at Harvard Univer- 
sity (MCZ). This collection represents a most important regional taxonomic resource, with 
superbly prepared and documented specimens, often with an abundance of affiliated host- 
plant and related biological information. The Winter material at the MCZ presently totals 
18,077 specimens, 16,079 from the 1996 donation and 1998 from donations between 
1979-1989. 

The strongest holdings are in the Noctuidae and Geometridae, although there is good 
representation of many groups. Virtually all the material was either collected personally 
by Dave, or reared by him from stock that he secured or obtained from other lepidopter- 
ists (few specimens are the result of direct exchange or gift). Dave ran a light trap for 
moths regularly in his back yard, with the back yard moving around in Westwood (from 
the early 1960s to 1975) and then on to nearby Dedham (1975 to 1995), in mostly semi- 
open environments with mixed hardwoods. Over time, the radius of his local butterfly and 
moth collecting expanded to include much of New England, Long Island, and northern 
New Jersey. Dave and his wife, Jo Brewer, also traveled extensively throughout North 
America to photograph and collect, with favored localities including Sanibel and Captiva 
Islands in Florida, and Ossibaw Island in Georgia. 

Table 1 provides a synopsis of the MCZ donation. Several hundred specimens remain 
with Dave at present. 


TABLE 1. Lepidoptera donated by William D. Winter to the Museum of Comparative 
Zoology at Harvard University. 


Year of Donation 


Group 1996 1979-1989 


Hesperiidae 827 374 
Papilionidae 436 33 
Pieridae 256 135 
Lycaenidae, Riodinidae 401 483 
Nymphalidae 762 110 
Satyridae Be 147 
Danaidae 10 6 
Pyralidae 140 = 
Thyatiridae, Drepanidae 92 — 
Geometridae 3052 — 
Mimallonidae, Apetalodidae, Lasiocampidae 94 = 
Saturniidae 389 — 
Sphingidae 296 — 
Notodontidae 542 = 
Arctiidae 786 — 
Lymantriidae 87 = 
Noctuidae 5443 810 
mixed families 1408 = 
exotics 837 a 
unsorted microlepidoptera 827 a 

16079 1998 


NAOMI PIERCE, Museum of Comparative Zoology, Harvard University, 26 Oxford 
Street, Cambridge, Massachusetts 02138, USA. 


Received and accepted for publication 15 January 1997. 


Journal of the Lepidopterists’ Society 
51(1), 1997, 103 


MANUSCRIPT REVIEWERS, 1996 


The merit of a scientific journal depends on the quality of its reviewers as well as its 
authors, but the former are usually unknown to readers. The Journal relied on the 
expertise of 61 reviewers last year to provide 108 evaluations of manuscripts. It is with 
much gratitude that the Journal acknowledges the services of the people listed below 
from whom manuscript reviews were received in 1996. Those who reviewed two or more 


manuscripts are denoted by asterisks. 


Annette Aiello, Balboa, Panama 
John Alcock, Tempe, AZ 
Richard A. Arnold, Pleasant Hill, CA 
“George T. Austin, Las Vegas, NV 
*Greg Ballmer, Riverside, CA 
*Carol Boggs, Stanford, CA 
*M. Deane Bowers, Boulder, CO 
Andrew V. Z. Brower, New York, NY 
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Donald Davis, Washington, DC 
Robert Dirig, Ithaca, NY 
Julian Donahue, Los Angeles, CA 
Boyce Drummond, Florissant, CO 
Thomas D. Ejichlin, Sacramento, CA 
“Terrance Fitzgerald, Cortland, NY 


Timothy P. Friedlander, North Potomac, 


MD 
David Gibo, Ontario, Canada 
David C. Hawks, Riverside, CA 
Martin Honey, London, England 
Ronald Hodges, Washington, DC 
Richard Holland, Albuquerque, NM 
Ian Kitching, London, England 
Ed Knudson, Houston, TX 
*Gerardo Lamas, Lima, Peru 
C. Don MacNeill, San Francisco, CA 
*Tim McCabe, Albany, NY 
*Eric H. Metzler, Columbus, OH 


Olaf Mielke, Parana, Brazil 
*William E. Miller, Saint Paul, MN 
*James Miller, New York, NY 

Michael Montgomery, Hamden, CT 
*John W. Peacock, Marion, OH 

Olle Pellmyr, Nashville, TN 

Kenelm Philip, Fairbanks, AK 
*Richard Peigler, Denver, CO 

Adam Porter, Bowling Green, OH 

John Rawlins, Pittsburgh, PA 
*Jim Reynolds, London, England 

Susan Richardson, Davis, CA 
*Robert K. Robbins, Washington, DC 
“Ronald Rutowski, Tempe, AZ 
*Phillip J. Schappert, North York, 

Ontario 
*James A. Scott, Lakewood, CO 

J. Marc Scriber, East Lansing, MI 
*Arthur M. Shapiro, Davis, CA 
*John Shuey, Indianapolis, IN 

Scott Smedley, Ithaca, NY 
*Felix A. H. Sperling, Berkeley, CA 
*Nancy Stamp, Binghamton, NY 

Ray Stanford, Denver, CO 

V. Thiagarajan, Coonoor, India 

Anthony Thomas, Fredericton, 

New Brunswick 

James Tumlinson, Gainesville, FL 
*David Wagner, Storrs, CT 

Ward B. Watt, Stanford, CA 
*Christer Wiklund, Lund, Sweden 

Ernest Williams, Clinton, NY 


Date of Issue (Vol. 51, No. 1): 4 April 1997 


EDITORIAL STAFF OF THE JOURNAL 


Lawrence F. Gat, Editor 
Computer Systems Office 
Peabody Museum of Natural History 
Yale University 
New Haven, Connecticut 06511, U.S.A. 
email: lawrence.gall@yale.edu 


Associate Editors: 
M. Deane Bowers (USA), Gerarpo Lamas (Peru), 
KeneLM W. Puiuie (USA), Rosert K. Rossins (USA), Fevix A. H. Speriinc (USA), 
Davip L. Wacner (USA), CuristeR WikLUND (Sweden) 


NOTICE TO CONTRIBUTORS 


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HEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp. 
1961a. Some contributions to population genetics resulting from the study of the Lepi- 
: doptera. Adv. Genet. 10:165-216. 


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CONTENTS 


PRESIDENTIAL ADDRESS 1996: ON THE BEAUTIES, USES, VARIATION, AND HAN- 
DLING OF GENITALIA John M. Burns 


DIVERSITY AND EVOLUTION OF TONGUE LENGTH IN HAWKMOTHS (SPHINGIDAE) 
William E. Miller 


THE IDENTITY OF FILATIMA ORNATIFIMBRIELLA (CLEMENS 1864) (GELECHIO- 
IDEA: GELECHUDAE) Ronald W. Hodges and D. Adamski 


AN EXAMPLE OF CLINAL VARIATION IN EASTERN NortH AMERICAN BUCKMOTHS 
(SaTURNUDAE: Hemiteuca) Brian G. Scholtens and Warren Herb 
Wagner fri re 


A NEW SPECIES OF RIODINIDAE FROM CoLomBia Curtis J. Callaghan and 
Julian Salazar 


A REVISION OF THE EUSELASIA ORFITA COMPLEX (RIODINIDAE) Curtis d- 


Callaghan 


TERMINATION OF PUPAL DIAPAUSE IN CALLOPHRYS SHERIDANII (LYCAENIDAE) 
Kiyoshi Hiruma, Jonathan P. Pelham and Hervé Bouhin 


NoTES ON THE MALE GENITALIA OF THE ANAEA RYPHEA - ANAEA EURYPYLE 
COMPLEX (NyMPHALIDAE) Astrid Caldas 


GENERAL NOTES 


Body weight as related to wing measure in hawkmoths (Sphingidae) William E. 
Miller 


Occurrence of the Palaearctic moth, Cnephasia longana (Tortricidae), on Santa 
Rosa Island, California J. A. Powell 


Effects of gene-environment interaction on silk yield in Antheraea mylitta (Saturniidae) 
A. K. Sengupta and A. A. Siddiqui. 0 


Distribution of a northern fauna of Noctuidae in the mountains of Oregon Paul C. 
Hammond and David:G. Grimble i 


W. D. Winter Collection to the Museum of Comparative Zoology, Cambridge Naomi 
Pierce. a ON Te 


MANUSCRIPT REVIEWERS, 1996 20020 ee 


32 


47 


57 


62 


75 


83 


This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). 


a 
>Volume 51 1997 ! Number 2 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 


Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS 


EMIT HSONTAY 


‘UG 01 1907, } 
LIBRARIES 


11 July 1997 


Ss 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE CouNCIL 


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President Vice President 

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Hon. Miriam Rotuscuiitp (1991), CLaupe Lemaire (1992) 


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Cover illustration: adult Catocala relicta Walker with the Peabody Museum of Natural. History at 


Yale University as backdrop, the logo for the 50th Anniversary Meeting of the Lepidopterists’ Society 


in July 1997. Original pen and ink drawing by Amy Bartlett Wright, Scientific Illustration, 202 Cot- 
tontail Drive, Portsmouth, Rhode Island 02871 USA. 


pe, Se ee a ee 


Mio 
a, 
os 
a a - 


7 


JoURNAL OF 
Tue LerpiporTeRIstTs’: SOCIETY 


Volume 51 1997 Number 2 


Journal of the Lepidopterists’ Society 
51(2), 1997, 105-118 


THE IMMATURE STAGES OF OXYTENIS MODESTIA, WITH 
COMMENTS ON THE LARVAE OF ASTHENIDIA AND 
HOMOEOPTERYX (SATURNIIDAE: OXYTENINAE) 


ANNETTE AIELLO 


Smithsonian Tropical Research Institute, P. O. Box 2072, Balboa, Ancon, 
Republic of Panama 


AND 


MANUEL A. BALCAZAR L.! 


Florida State Collection of Arthropods, Division of Plant Industry, Florida 
Department of Agriculture and Consumer Services, P. O. Box 147100, 
Gainesville, Florida 32614, USA 


ABSTRACT. The immature stages of Oxytenis modestia are described, with special 
attention to the first instar, the larval food plants are reported, and larval and adult behav- 
iors are described. Limited information is given for the genera Homoeopteryx and As- 
thenidia, and they are compared and contrasted with Oxytenis. 


Additional key words: Life history, Alibertia edulis, Genipa americana, Rubiaceae. 


The genus Oxytenis Hiibner includes 17 described species of 
medium-sized moths (Jordan 1924) that are patterned to resemble 
dried, brown leaves (Fig. lc). Oxytenis and two other genera, Astheni- 
dia Westwood and Homoeopteryx Jordan, have been treated as the fam- 
ily Oxytenidae (Jordan 1924). Minet (1994) showed that these three 
genera do form a monophyletic group and, because he considered them 
to represent the most ‘primitive’ lineage of the Saturniidae, he reas- 
signed them to that family as the subfamily Oxyteninae. 

Unfortunately, technical descriptions for the larvae of Oxyteninae are 
still lacking. Brief, superficial descriptions are available for the larvae of 
some species: Oxytenis naemia Druce, O. angulata (Cramer), O. ferru- 
ginea Walker, O. modestia (Cramer) and Asthenidia lactucina (Cramer) 
(Jordan 1924, Draudt 1929). Nentwig (1985) described and illustrated 


! Present address: Departamento de Zoologia, Instituto de Biologia, Universidad Nacional Au- 
tonoma México, Apdo. Postal 70-153, 04510 Mexico D. F.. MEXICO 


106 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the striking sequence of mimicry presented by various instars of O. 
naemia. Miles Moss produced a series of beautiful water color paintings 
of various oxytenine larvae, but these were never published and are in 
the Natural History Museum, London. Here we describe the immature 
stages and larval biology of Oxytenis modestia. 


MATERIALS AND METHODS 


Oxytenis modestia (Fig. 1) was reared by the senior author on several 
occasions, from larvae collected on Alibertia edulis (A. Rich.) A. Rich. 
and Genipa americana L. (Rubiaceae) (see Table 1 for list). The only 
eggs obtained (Aiello Lot 80-9) were from a female collected at light on 
Barro Colorado Island, Panama, on 8 March 1980. 

Of the 96 eggs produced by the female, ten were preserved in 70% 
ethanol. The remaining 86 eggs were divided into four groups, placed in 
petri dishes, each with a piece of moist folded paper towel, and kept in 
ZipLoc bags in an air conditioned laboratory. Upon hatching, the larvae 
were offered squares of 11 different species of plants, all members of 
the Rubiaceae: Alibertia edulis, Alseis blackiana Hemsl., Coussarea 
curvigemmia Dwyer, Faramea occidentalis (L.) A. Rich., Hamelia patens 
Jacq., Isertia haenkeana DC., Ixora coccinea L., Palicourea guianensis 
Aubl., Psychotria grandis Sw., Psychotria marginata Sw., and Randia ar- 
mata (Sw.) DC. The larvae preferred young leaves of Alibertia but ate 
small amounts of young Alseis, Ixora, and Randia as well. They were 
reared on Alibertia edulis. As they developed, the majority of the 86 in- 
dividuals were preserved at various stages as vouchers, and only 12 indi- 
viduals were reared to adults. 

Specimens reared before 1981 are at the National Museum of Natural 
History (NMNH), Washington DC; those reared from 1981 to the pres- 
ent are at the Smithsonian Tropical Research Institute (STRI), Panama. 
Voucher specimens of all instars reared from eggs were preserved in 1:1 
xylene : ethanol-95% (XA) and are at the NMNH. To study the morphol- 
ogy of the first instar in detail, two larvae were prepared for viewing in a 
Scanning Electron Microscope (SEM). The larvae were dehydrated in 
graded series of ethanol (70%, 80%, 90%, and 100%), then critical point 
dried in a DCP-1 Denton Vacuum with carbon dioxide. Specimens were 
then mounted on points and coated with a Denton Vacuum Desk II 
sputter coater, using a gold target, for 6 minutes (15 Ma). Prepared 
specimens were held on micro test clips (Radio Shack 270-373) and 
mounted on stubs for observation with a Hitachi 570s (15kV) SEM. 

Morphological terms follow Dethier (1941) for the antennae, Grimes 
and Neunzig (1986a, 1986b) for the maxillae, Beck (1961) and Miller 
(1991) for tarsal setae, Stehr (1987) for chaetotaxy and general morphol- 
ogy, and Mosher (1916) for the pupae. 


VOLUME 51, NUMBER 2 107 


(RRS asa than hs Pics. Meg oe ee) Serene Meer V LES - a fase 


Fics 1-7. 1-4, Oxytenis modestia: 1 (upper): adult male (Aiello Lot 85-26), 1 (mid- 
dle): adult fale (Aiello Lot 81-65), 1 (lower): adult female (natural position) (Aiello 
Lot 84-55 no.4), 6.5 cm across; 2, final instar (green morph) (Aiello Lot 80-9 no.1), 3.6 cm 
long; 3, final instar (brown morph) (Aiello Lot 87-52 no.1); 4, fourth instar (Aiello Lot 
87-63), 2.6 cm long. 5, bird-dropping, 2.1 cm long. 6, Homoeopteryx malecena early final 
instar (Aiello Lot 79-122), 3.1 cm long. 7, Asthenidia transversaria final instar (Aiello Lot 
81-37), 3.5 cm long. 


108 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


IMMATURE STAGES 


Stock source. The female moth, collected as Aiello Lot 80-9, on 8 March, began 
ovipositing shortly after being placed in a wire and petri dish cage with a leaf of Faramea 
occidentalis. Before dying on 12 March, she laid 96 eggs on the floor paper and the wire 
walls, as well as on the leaf. 

Egg. Duration 5 days. Ellipsoid, approximately 1.3 mm in length, green. The eggs were 
green when fresh, but gradually faded to yellow by day five. By day three, the stemmata 
began to show through the chorion of some individuals, and by day four the mandibles 
were visible. On day five, the final day of the egg stage, the larvae could be seen clearly. 
They were folded twice to form a condensed ‘S,’ and they slowly shifted position within 
the egg. 

First instar, general. Duration 4—5 days. Pale beige in color; head 0.6 mm wide; bod 
3.8—4.2 mm long; primary setae represented by granulate setae and chalazae with bulbous 
glandular tips that secrete a sticky substance. Head: (Figs 8, 9), hypognathous; surface 
smooth with long primary setae and without secondary setae, F1 very short, C2 arises from 
the lateral margin of the clypeus and runs parallel to the head surface, Al shortest A seta, 
S2 and A3 well above the stemmatal semicircle; stemmata 1—2 and 3—4 very close forming 
a semicircle, stemma 5 distant and ventral to 6; ecdysial line and fronto-clypeal suture in- 
conspicuous; adfrontal suture conspicuous only for upper half of the frons; anteclypeus 
grooved; labrum with six pairs of setae, without notch; mandibles quadrate with five op- 
posable teeth; antennae (Fig. 11) prominent, segment 1 short; segment 2 with two sensilla 
trichodea (ST) on the external side, sensillum anterior several times larger than caudal 
sensillum, with three sensilla basiconica (SB), the larger ones near the anterior and poste- 
rior margins of the segment, the small SB half-way between the anterior SB and ST; seg- 
ment 3 projecting, located medially with respect to $2; sensillum styloconicum (SC) as 
long as longest sensillum basiconicum, and located in the posterior and anterior margins 
respectively, two small SB medially and laterally; maxillary lobe with STI removed ventro- 
laterally from and larger than STIIJ—III, all acicular; MSS and LSS subequal; all other sen- 
silla inconspicuous; maxillary palpus with all eight sensilla basiconica subequal; spinneret 
(Fig. 12) dorsoventrally compressed, truncate. Thorax: relatively smooth; spiracle oval, as 
large as that on A7; primary setae XD, D and SD represented by chalazae with setae bear- 
ing bulbous glandular tips (Figs. 9, 10, 16), except for D1 and D2 on T1 and D2 on T2, 
which are thick setae; L group with two setae on T1, and one on T2—3; two SV and one V 
on all three segments; thoracic legs with seta 2 (Ts2) acicular (Fig. 13), Ts3 and Ts] lance- 
olate, Ts3 slightly longer than Ts1. Abdomen: A1—8 with D1 and SD1 as chalazae with 
gland-tipped setae, only one SD, L, and SV setae; a minute, round spiracle on Al—6, A7 
with an oval larger spiracle, A8 with largest, circular spiracle; a chalaza with gland-tipped 
seta as D1 on anal shield. Crochets 6 or 7, in a uniordinal homoideous mesoseries plus 4—5 
crochet remnants of a uniordinal lateroseries. Crochets not deeply set in the spatula on 
the proleg; the fleshy lobe of the proleg gives the crochets a more or less C-shaped pat- 
tern (Fig. 15). The first instars emerged from their eggs the morning of day six and, after 
testing them with a variety of rubiaceous plants, all were reared on Alibertia edulis. 

Second instar. Duration 3—5 days. Similar to first instar except that the chalazae are 
larger and are drawn out to form a stalk, tipped by a sticky gland; the metathoracic, L cha- 
laza is located on a tiny triangular lateral flange and is gland-tipped; the D1 chalazae of A8 
are located at the tip of a fleshy ‘tail,’ and the larva is now darker and has the overall ap- 
pearance of a bird dropping. 

Third instar. Duration 4—6 days. A larger version of the second instar, except that it is 
black with large brown chalazae (Fig. 4) and is an excellent mimic of a bird dropping filled 
with seeds (Fig. 5). 

Fourth instar. Duration 3—5 days. Essentially the same as the third, only larger. 

Fifth instar. Duration 6—7 days if final, 3-4 days if not final. This species has a vari- 
able number of stadia, independent of sex, and the final instar is different from the others, 
whether instar 5 or 6. If the fifth instar is not the final instar, it looks essentially the same 
as the three previous instars, only larger. If the fifth instar is the final instar, it takes on a 
different appearance, a description of which appears under “sixth instar.” 


VOLUME 51, NUMBER 2 


TABLE 1. 


reared by the senior author. 


109 


Oxytenis modestia, Homeopteryx macelena, and Asthenidia transversaria 


Stage Number adults Larval food plant Aiello 

collected obtained (Rubiaceae) lot # 
Oxytenis modestia 
Eggs feome Df 80-9 
Third to final instar ee Genipa americana L. 84-55 
Second to final instar lm Genipa americana L. 85-126 
Penultimate instar lm Alibertia edulis (A. Rich.) A. Rich.in DC. 81-43 
Penultimate instar ese Alibertia edulis (A. Rich.) A. Rich.in DC. 81-65 
Penultimate instar lm Genipa americana L. 87-52 
Final instar 1h Alibertia edulis (A. Rich.) A. Rich.in DC. 80-46 
Final instar ie Genipa americana L. 78-90 
Pupa ise Genipa americana L. 78-91 
Homoeopteryx malecena 
Penultimate instar iat Faramea occidentalis (L.) A. Rich. 82-24 
Second to final instar Lee Faramea occidentalis (L.) A. Rich. 82-49 
Third to final instar lm Faramea occidentalis (L.) A. Rich. 82-78 
Penultimate instar 1b Faramea occidentalis (L.) A. Rich. 79-122 
Final instar ie Faramea occidentalis (L.) A. Rich. 80-74 
Asthenidia transversaria 
Final instar 1-Sé Calycophyllum candidissimum (Vahl) DC. 81-37 
Penultimate instar lm Warscewiczia coccinea (Vahl) K1. 81-67 


Sixth instar: Duration 6—8 days. Regardless of whether it is the fifth or the sixth, the 
final instar looks quite different from any of the preceding instars and it occurs as either a 
green or a brown morph, with variations. In all cases, the metathorax bears subdorsal eye 
spots (black, bordered by yellow) towards the anterior margin. These ‘eyes’ are located in 
a fold of cuticle and thus can be ‘opened’ and ‘closed’ by raising or lowering blood pres- 
sure to the thorax. At rest the eyes are closed, but a highly disturbed larva inflates the tho- 
rax, thus opening the ‘eyes,’ and rears up, waving back and forth like an angry serpent. In 
this stadium, the glands are tiny and sessile on the body, and the L gland of the metatho- 
rax is missing. The green morph (Fig. 2) is dark green with a bright green dorsal stripe that 
widens on A4, narrows on A5, then widens again to cover the sides of the body on A6-8. 
A long black dorsal diamond mark on A6—8 terminates with the ‘tail.’ Occasional individu- 
als have the dorsal stripe cryptically marked with brown, yellow, green, and black. Brown 
individuals (Fig. 3) are like the green ones except that the ground color and dorsal stripe 
are brown; the head remains green, however. 

Larvae, general: Individuals passed through five to six larval instars (Table 2). Larval 
stadia were trimorphic, that is the general appearance of each larva changed abruptly 
upon the molt to second instar and changed again upon molting to the final instar, regard- 
less of whether that final instar was the fifth or the sixth. The condition of trimorphic lar- 
val stadia together with a variable number of larval instars occurs as well in the nymphalid 
Dynastor darius (Fabricius) (Aiello, 1978), and in the saturniid Arsenura batesii (Aiello, 
pers. obs.). In the case of Oxytenis the situation is more complex because, in addition, the 
larvae are polymorphic for color pattern within the later intermediate stadia and the final 
stadium; some individuals are green (Fig. 2) and some are brown (Fig. 3), regardless of 
sex, and they show varying degrees of pattern complexity. As the result of this variation in 
color and development the larger larvae of a brood tend not to resemble one another; the 
multiple search images resulting from such variation perhaps make them less easy for a 
predator to locate. 

From the outset, the larvae assumed a ‘J’ position, with the head and thorax turned back 


110 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ie : te. 


Fics. 8-11. 8, Oxytenis modestia first instar, head (front view) (Aiello Lot 80-9). A = 
anterior setae, AF = adfrontal setae, C = clypeal setae, F = frontal setae, L = lateral setae, 
P = posteriodorsal setae, S = stemmatal setae. Scale line = 0.15 mm; 9, Oxytenis modestia 
first instar, head and thorax (lateral view) (Aiello Lot 80-9). Scale line = 0.3 mm; 
10, Oxytenis modestia first instar, abdominal segments 6—10 (lateral view) (Aiello Lot 80- 
9). Scale line = 0.25 mm; 11, Oxytenis modestia first instar, right antenna (ventral view, 
with anterior at top) (Aiello Lot 80-9). Scale line = 11.55 um. 


to one side and pressed against the abdomen when mildly disturbed or when at rest, a be- 
havior that continued throughout larval life, and which, especially in the intermediate in- 
stars, gave the larva the appearance of a bird dropping. All larval instars bore sticky glands. 
Perhaps to avoid contamination of these glands, the larvae grasped their fecal pellets in 


VOLUME 51, NUMBER 2 111 


Fics. 12-15. 12, Oxytenis modestia first instar, hypopharingeal complex (dorsal view 
with anterior at top). (Aiello Lot 80-9). Scale line = 37.5 um; 13, Oxytenis modestia first 
instar, right pretarsus of mesothoracic leg (mesal view) (Aiello Lot 80-9). Scale line = 25 
um; 14, Oxytenis modestia first instar, gland-tipped setae (D on A2—3, detail) (Aiello Lot 
80-9). Scale line = 37.5 um; 15, Oxytenis modestia first instar, left proleg (Aiello Lot 80- 
9). Scale line = 25 um. 


their mandibles as they were produced and dropped them off the leaf. Interestingly, the 
earliest instars of various Oxytenis species are superficially similar, in color and pattern, to 
the earliest instars of various species of Adelpha Hiibner (Nymphalidae) that feed on the 
same array of rubiaceous plants (Aiello, pers. obs.). The larvae remained on the upper sur- 
face of their food leaves except during molting, at which time they moved to the wall of 


112 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


5 
5 


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<2) 
as Ea a AA GAA AO, A A 
Bx Ay Ay Ay Ay Ay Ay Ay A Oy Ay Ay 
ac PAA AAA AAA A AA, 
= A, Ay Ay A, A, Ay A, Ay Ay AY 
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fifth instar, 6 = sixth instar, P 


dividuals and sixth for wee 


fourth instar, 5 


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Individual 


TABLE 2. Development times of the twelve Oxytenis modestia reared as Aiello Lot 80-9; e = egg, 1 
Rie ee ec e lid 2 2 293393) 34 4944 525) 5 626766, 6-6 P PoP PR Pee PP PP BP female 


lIeverene el o  l 222 Oe Sa ee oa oa 444 be ae > 

Boeveve 6.6 ttle dso. Be Oe a 4d 4A Son > 

4e e ee e@ lod 1.2 2. Beer a OA a 4 6566 6 86N6 652° PYPePePeeS Ee. 
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S eme-e €e-F bk tk bo 24622 SoS - 8. Set 44425 3555 6:6 646 -656-k2) P 
Geeee el Lol tf Fe 2-2 2 2 273) 3°3.303- 44 4.475 525 5 676) 6 

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Ise eee el Pa 1 ao 2S Boe 824-474 45555) 5.6 66.6 Gu6 


instar, 4 
Day 


VOLUME 51, NUMBER 2 1S 


SD2, 


SAN 
ey a a 
16 pe Yee» 


WZ 13 


Fic. 16. Oxytenis modestia first instar, setal map. A = abdominal segment, D = dorsal 
setae, L = lateral setae, s = spiracle, SD = subdorsal setae, SV = subventral setae, V = ven- 
tral seta, XD = extradorsal setae. 


the petri dish. In the wild, larvae of Oxytenis modestia usually move to the petioles and 
branches to molt, although they may remain on the blade (Aiello, pers. obs.). 

Pre-pupa: Much faded and shortened, with the ‘eyes’ opened. Mature larvae con- 
structed very loose cocoons of leaves and wet, stretchy, reddish silk on the floor of the 
cage, and they produced a copious puddle of liquid in which the pre-pupae rested. Being 
shrunken, faded, and wet, they appeared dead and rotten, but, the next morning, they pu- 
pated. The production of liquid by pre-pupae has been reported for two sphingids, Man- 
duca (as Protoparce) rustica (Fabricius) (Moss 1912) and Xylophanes mossi Clark (Moss 
1920). 

Cocoon: Very loose cocoon of leaves and wet, stretchy, reddish silk. 

Pupa (Fig. 17a shows Oxytenis naemia): Duration 12—14 days. Obtect; fronto-clypeal 
suture obsolete; labial palpi not visible; pilifer lobes absent; the maxillae and prothoracic 
legs the only appendages visible within the area delineated by the antennae; antennae 
broadly pectinate, touching for the apical 0.5—2 mm, the width one seventh the length; 
mesothoracic wings, on the ventral surface of the body at meson, shorter than caudal mar- 
gin of the fourth abdominal segment; abdominal segments with scattered minute setae not 
visible without magnification; cremaster with approximately 24 reddish hooks, which are 
quite tangled together. 

Larval food plants:. Alibertia edulis and Genipa americana (Rubiaceae). 

Adults. Adults eclosed around 11 pm and the females began emitting pheromone 
about 2.5 hours later. The resemblance of the adults to dried, brown leaves is greatly en- 
hanced by the behavior of the moths, which rest with the wings held so as to form a ‘leaf, 
complete with midrib, petiole, and drip tip. When disturbed, the moths release their grip 
on the substrate and gently waft to the ground. 

Distribution and flight period. Guatemala to Bolivia and southeastern Brazil, pre- 
sumably occurring also in Mexico and northern Argentina; flying during March to Decem- 
ber (Jordan, 1924). On Barro Colorado Island, adults can be seen at any time of year, al- 
though they are rare from November through April, and are most abundant from May 


through July. 


NOTES ON HOMOEOPTERYX AND ASTHENIDIA 


Homoeopteryx malecena Druce has been reared by the senior author 
on three occasions from larvae found on Faramea occidentalis, and As- 
thenidia transversaria Druce has been reared twice from larvae, once on 
Calycophyllum candidissimum (Vahl) DC. and once on Warscewiczia 
coccinea (Vahl) KI. (Table 1). All three plants belong to the Rubiaceae. 


114 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 3. Distribution of sticky glands for Homeopteryx, Asthenidia, and Oxytenis. 
Values in table are total number of sticky glands per segment, for thoracic segment | 
through abdominal segment 10. 


Tl T2 T3 Al A2 A3 A4 A5 A6 AZT A8 AQ Al1O 


Homoeopteryx early instars 8 8 8 4°°4 <4) 4. 4y A AeA eee 
final instar 8: 10 10 .6°.6,% 66 - 66 “GR SGeaeiGweG 
Asthenidia final instar 8 6 102.4 4° “4. 4) 4 4A Ao 
Oxytenis early instars 8 6. (8. 4 4.4, 4: 2A ae 
final instar 8 (6.10 4 4.4.94. 4) 4A Ae 


Homoeopteryx. Although adults of Homoeopteryx are superficially 
very similar to those of Oxytenis in general appearance, the larvae of the 
two genera differ in several respects. Although all instars we have seen 
bear sticky glands and rest in a ‘]’ position, they lack the flanges and eye- 
spots of Oxytenis, and the gland pattern of Homoeopteryx appears to be 
more complete than that of Oxytenis (Table 3). 

The youngest larva seen (Aiello Lot 82-78), was cream colored along 
the dorsum and purple-brown along the sides. Its sticky glands were at 
the ends of long setae and were distributed as shown in Table 3 for early 
instars. The head had long white setae, without glands, and glandless se- 
tae were also found among the gland-tipped ones. The distribution of 
setae with glands was the same as for early instar Oxytenis except that 
Oxytenis had only 6 gland-tipped setae on T2, whereas Homoeopteryx 
had 8. Judging by the long, gland-tipped setae, it is most probable that 
this early larva was a first instar, and if so, then that individual had only 
four larval instars. 

The next instar was cream, cryptically patterned with black, and had a 
poorly-defined cream saddle on segments A4—A6. The glands were ses- 
sile on slender chalazae instead of at the tips of setae. The chalazae of 
T3 were located on an annular swelling, which could be folded forward 
against the body. The D chalazae (with sticky glands) of A8 were at the 
apex of a short ‘tail’ giving that protuberance a slightly forked appear- 
ance. The gland pattern changed (Table 3) in that T2 and T3 now had 
10 instead of 8 glands, and the abdominal segments now each had 6 
glands instead of 4. 

The following instar was a larger, paler version of the previous one, 
except that the T3 swelling was more pronounced and the ‘tail’ was 
rather thick. . 

The final instar was at first quite dark, although the cream saddle was 
still visible. As the larva matured, it became less dark, more evenly cryp- 
tically marked, and the saddle mark darkened somewhat. In general, the 
larva became an excellent twig mimic, a resemblance that was greatly 
enhanced by the T3 annular swelling, the slightly paler saddle mark, and 


VOLUME 51, NUMBER 2 tales 


Fic. 17. Pupae. a, Oxytenis naemia, ventral view (Lot 81-80 no.4), 2.4 cm long; b, As- 
thenidia transversaria, ventral view (Lot 82-67), 1.4 cm long; e-d, Homoeopteryx male- 
cena, ventral, lateral, and dorsal views (c: Lot 82-78, d: Lot 82-24), each 2.2 cm long. 


the fact that the larva rested with the body held rigid with the anterior 
end held up at a slight angle, in the manner of many geometrids. 

Among other individuals reared, the intermediate instars were gray 
and cryptically marked with a green and brown lichen-like pattern. Ma- 
ture larvae may also be cryptically marked or may be the same mossy 
green of their food plant leaves (Fig. 6). 

The larva made a leaf and silk cocoon in which it pupated several days 
later. The pupa (Fig. 17c,d), instead of the uniform dark brown to black 
of an Oxytenis was dark beige with black markings, and the pattern var- 
ied among individuals. 

Asthenidia. Adults of Asthenidia bear no resemblance to either 
Oxytenis or Homoeopteryx, and instead look like small, white swallow- 
tail butterflies. However, the ‘tail’ and eyespots of the mature larva 


116 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(Fig. 7) reveal it to be an oxytenid. The larva reared on Calycophyllum 
candidissimum, was nearly mature when collected and the one reared 
on Warscewiczia coccinea was collected the day before it molted to the 
larval final stadium. Both larvae rested in a ‘J’ position on the upper sur- 
face of leaves. 

The younger of the two larvae bore stalked glands with non-glandular 
setae at the bases. The glands were arranged as in the early instars of 
Homoeopteryx. There were false eyes on T3 that could open and close 
as in Oxytenis. Although there were no thoracic flanges, the metathorax 
(T3) was slightly expanded and each expansion bore a single gland. As 
well, there were two humps, each with two peaks, one on T3 and one on 
A3. The ‘tail’ on A8 had two glands at the apex and had an extra point 
posterior to and below the apex. The larva was cryptically patterned with 
gray, brown and whitish, and had a low lateral white area on A3—4 and 
A7-8. 

The final instar had deep blue eye spots with a broad black border. 
The thorax expansions were now almost non-existent. The hump on T3 
was high and had two peaks close together. A3 had two large bumps. 
The tail had a posterior bump at the mid point. The larva was very dark 
green to black, except dorsally from the posterior side of the T3 hump, 
narrowing between the A3 humps, to a small dorsal green triangle on 
A4, then widening on A5 to a broad green dorsal stripe that swept up 
and included the ‘tail.’ There was a subventral white mark on each of A4 
and A7—8. Many glands were missing and the few remaining glands 
were sessile and were best formed on segments T1—A8. 

The other final instar was similar but was brown instead of green, and 
the black was confined to the subventral areas and to bold oblique lat- 
eral marks on A3, A4, small lateral triangles on A6 and A7, and a dark 
lateral crescent on A4—6. It also had a subventral white mark on A7-—8. 
The only definite glands were on the prothorax, in two groups of four, 
and they were sessile. 

Both larvae made loose cocoons of leaf and silk. The pupae (Fig. 17b) 
were short compared with those of Oxytenis, and the abdomen came to 
a more acute point. 


SYSTEMATIC RELATIONSHIPS 


The possession of sticky glands by the larvae, the habit of resting in a 
‘| position, the utilization of rubiaceous food plants, and the similarities 
of pupal morphology seem to tie the three genera, Homoeopteryx, As- 
thenidia, and Oxytenis together. The distribution of sticky glands is sim- 
ilar for the three genera in the early stadia. In later stadia, the distri- 
bution is more complete for Homoeopteryx than for Oxytenis or 
Asthenidia (Table 3). That, plus the fact that Homoeopteryx has no tho- 


VOLUME 51, NUMBER 2 JEAN 


racic flanges or eye spots leads us to believe that that genus is more gen- 
eralized and nearer to the ancestral condition than are the other two. 
Asthenidia, with its eye spots and thoracic expansions is more reminis- 
cent of Oxytenis. Based upon a handful of rearings, Homoeopteryx and 
Asthenidia may each be confined to a more limited selection of larval 
food plant, while the various species of Oxytenis are found on plants of 
at least five genera, Alibertia, Faramea, Genipa, Isertia, and Posoqueria. 
Both Jordan (1924) and Michener (1952) indicated that the use of lar- 
val characters, especially from the first instars, in Oxyteninae and Cer- 
cophaninae could be of great help in understanding phylogenetic rela- 
tionships among Saturniidae. Nevertheless, the known larvae of 
Oxytenis show many specialized characters, such as the sticky scoli, that 
make it very difficult to determine character homology. A similar situa- 
tion was found for Cercophana venusta (Walker) (Wolfe & Balcazar 
1994). Nassig (1989). proposed a classification of the scoli found in the 
Saturniidae (not including the Oxytenidae), but the peculiar type found 
in the Oxyteninae does not conform to any proposed classification. In- 
terestingly, Oxyteninae and Cercophaninae, the two most plesiomorphic 
subfamilies in Minet’s classification, have the D scoli on A8 not fused on 
the dorsomeson, the most characteristic feature of the bombycoid lar- 
vae that have few secondary hairs (“naked”) (Lampe & Nissig 1989, 
Nassig 1994, Oberprieler & Duke 1994). Fused D scoli on A8 was re- 
garded as a synapomorphy for the Bombycoidea by Minet (1994), under 
the asumption that its absence in some taxa (bombycoid larvae clothed 
in long secondary hairs, “wooly,” and the “naked” larvae of Salassa 
Moore [Salassinae], Anisota Hiibner, and Dryocampa Harris [Cerato- 
campinae]) is the result of secondary loss. It remains to be resolved 
whether these structures are synapomorphic or the result of conver- 
gence reflecting the same groundplan (cf. Oberprieler & Duke 1994). 


ACKNOWLEDGMENTS 


We are grateful to the Entomology and Nematology Department, University of Florida 
for the use of their scanning electron microscope, and to the STRI Electronic Imaging 
Laboratory for preparing the color figures. Many thanks are due Marshall Hasbrouck for 
preparing the illustrations of the pupae. Thanks also to David Hawks and two anonymous 
reviewers for their extremely helpful comments. 


LITERATURE CITED 


AIELLO, A. 1978. Life history of Dynastor darius (Lepidoptera: Nymphalidae: Brassoli- 
nae) in Panama. Psyche 85:331—345. 

BECK, H. 1960. Die Larvalsystematik der Eulen (Noctuidae). Nr. 4. Berlin: Akademie- 
Verlag. 

ee y. G. 1941. The antennae of lepidopterous larvae. Bull. Mus. Comp. Zool. 
87:455—507. 

DRAUDT, M. 1929-1930. Saturnidae [sic]. In A. Seitz (ed.), Die Gross-Schmetterlinge der 
Erde. A. Kernen, Stuttgart. pp. 713-827, Pls Olas A: 


118 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


GRIMES, L. R. & H. H. NEUNZIG. 1986a. Morphological survey of the maxillae in last stage 
_ larvae of the suborder Ditrysia (Lepidoptera): palpi. Ann. Entomol. Soc. Am. 
79:491—509. 
———. 1986b. Morphological survey of the maxillae in last stage larvae of the suborder 
| Ditrysia (Lepidoptera): mesal lobes (laciniogaleae). Ann. Entomol. Soc. Am. 
—79:510—526. 

JORDAN, K. 1924. On the Saturnoidean families Oxytenidae and Cercophanidae. Novit. 
Zool. 31:135—193, pls. 6-21. 

LAMPE, R. E. J. & W. A. Nassic. 1989. Neue Erkenntnisse uber die Gattung Lemaireia: 
1. Beschreibung der Praimaginalstadien von L. luteopeplus aureopeplus Nassig & 
Holloway (Lepidoptera, Saturniidae). Nachr. entomol. Ver. Apollo 10:225—231. 

MICHENER, C. D. 1952. The Saturmiidae (Lepidoptera) of the western hemisphere. Mor- 
phology, phylogeny, and classification. Bull. Am. Mus. Nat. Hist. 98:335—501, pl. 5. 

MILLER, J. S. 1991. Cladistics and classification of the Notodontidae (Lepidoptera: Noc- 
tuoidea) based on larval and adult morphology. Bull. Am. Mus. Nat. Hist. 204:1—230. 

MINET, J. 1994. The Bombycoidea: phylogeny and higher classification (Lepidoptera: 
Glossata). Ent. Scand. 25:63—88. 

MOSHER, E. 1916. A classification of the Lepidoptera based on characters of the pupa. 
Bull. Ilinois State Lab. Nat. Hist. 12:13—159 + pls 19-27. 

Nassic, W. A. 1989. Wehrorgane und Wehrmechanismen bei Saturniidenraupen (Lepi- 
doptera, Saturniidae). Verhand. Westdeutscher Entomologentag 1988:253—264. 

. 1994. Vorschlag fiir ein neues Konzept der Gattung Saturnia Schrank 1802 (Lep- 
idoptera: Saturniidae). Nachr. Ent. Ver. Apollo 15:253—266. 

NENTWIG, W. 1985. A tropical caterpillar that mimics faeces, leaves and a snake (Lepi- 
doptera: Oxytenis naemia). J. Res. Lepid. 24:136-141. 

OBERPRIELER, R. G. & N. J. DUKE. 1994. The life history and immature stages of Spi- 
ramiopsis comma Hampson, 1901 (Lepidoptera: Bombycoidea), with comments on its 
taxonomic position and on preimaginal characters of the Bombycoidea. Nachrichten 
Ent. Ver. Apollo 15:199—244. 

STEHR, F. W. 1987. Order Lepidoptera, pp. 288-305. In F. W. Stehr (ed.), Immature in- 
sects. Kendall/Hunt, Dubuque. 

WOLFE, K.' L. & M. A. BALCAZAR-LARA. 1994. Chile’s Cercophana venusta and its imma- 
ture stages (Lepidoptera: Cercophanidae). Trop. Lepid. 5:35—42. 


Received for publication 18 December 1995; revised and accepted 2 May 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 119-127 


A NEW SPECIES OF PHANETA, WITH TAXONOMIC 
DIAGNOSES AND SEASONAL AND GEOGRAPHICAL DATA 
ON FOUR RELATED SPECIES (TORTRICIDAE) 


DONALD J. WRIGHT 
3349 Morrison Avenue, Cincinnati, Ohio 45220, USA 


RICHARD L. BROWN 


Mississippi Entomological Museum, Box 9775, Mississippi State, 
Mississippi 39762, USA 


AND 


LORAN D. GIBSON 
8496 Pheasant Drive, Florence, Kentucky 41042, USA 


ABSTRACT. Phaneta canusana, new species, is described and compared with P. 
lapidana, P. sublapidana, P. kokana and P. ambodaidaleia. Lectotypes are designated for P. 
lapidana and P. sublapidana. Imagos of the five species, the male genitalia of P. lapidana 
and P. sublapidana and the male and female genitalia of P. canusana are illustrated. P. 
canusana is associated with prairie remnants in Ohio, Kentucky, Missouri, and Mississippi. 
New distribution records are given for P. kokana and P. ambodaidaleia. 


Additional key words: vernal flight, autumnal flight. 


The genus Phaneta Stephens consists of 102 species in North Amer- 
ica, including eight described since publication of the most recent check 
list for North American members of the genus (Powell 1983). This 
group was treated by Heinrich (1923) as Thiodia, distinguishable from 
Eucosma by the absence of a costal fold on the male forewing. Obraz- 
tsov (1952) restricted Thiodia to a group of European species and con- 
sidered Phaneta to be the correct generic name for the Nearctic species. 

While surveying the lepidopteran fauna of Lynx Prairie Preserve, 
Adams County, Ohio, in 1989 and the Osborn prairie remnant of the 
Mississippi Black Belt in 1991, we recorded an unknown species of 
Phaneta, described below as new. Representatives of this same species 
had been collected in Missouri by J. Richard Heitzman in 1976, and 
they were brought to our attention by W. E. Miller, who recognized the 
conspecificity of the Missouri and Ohio specimens. The new species has 
similarities with four other members of the genus: P. lapidana (Walsing- 
ham), P. swblapidana (Walsingham), P. kokana (Kearfott), and P. ambo- 
daidaleia Miller. 

In examining olethreutine type specimens residing in European col- 
lections, Obraztsov selected specimens to serve as lectotypes for Wal- 
singham’s species, but he never published those designations. His notes 
and photographs of this material are currently on loan to the Mississippi 


120 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Entomological Museum from the American Museum of Natural History 
(AMNH), and based on that information, lectotypes are designated in 
this paper for P. lapidana and P. sublapidana. | 

During this study, specimens from the following institutional and per- 
sonal collections were examined: Horatio T. Enterline (HTE), Loran D. 
Gibson (LDG), J. Richard Heitzman (JRH), Illinois Natural History 
Survey (INHS), Mississippi Entomological Museum (MEM), United 
States National Museum of Natural History (USNM), Donald J. Wright 
(DJW), and Zoological Museum, Copenhagen (ZM). 


Phaneta sublapidana (Walsingham) 
(Figs. 1, 2) 


Semasia sublapidana, Walsingham 1879:59, pl. 73, fig. 9 (imago) 

Thiodia sublapidana, Fernald [1903]:462; Heinrich 1923:50, fig. 122 (male genitalia); Mc- 
Dunnough 1939:44. 

Eucosma sublapidana, Barnes & McDunnough 1917:no. 7086. 

Phaneta sublapidana, Powell 1983:33. 


Semasia sublapidana was described from four males collected near Klamath Lake, Ore- 
gon. A lectotype for Semasia sublapidana Walsingham is hereby designated (Fig. 1): 
male, “(nr. Fort Klamath), Jackson Co., Oregon, 21—23 IX 1871, Wlsm.” BM genitalia slide 
11599 (Fig. 2). The lectotype is deposited in The Natural History Museum, London 
(BMNH). According to Obraztsov’s notes on the envelope containing the photograph of 
the lectotype, this was the specimen figured by Walsingham (1879). 

We examined the following specimens: California: Mono Co., NE side Mono Lake, 
6500', XI-19-93, D. Giuliana, 1400 PST [Pacific Standard Time], 46° F (1). Inyo Co., 
Deep Spr. Vy., 5300', XI-23-93, D. Giuliana, 1600 PST, 36° F (1) (UCB). J. Powell (pers. 
comm.) collected a series of 18 males during the second week of December, 1995 in the 
Owens Valley, Inyo County. These individuals were flying just before sunset, which occurs 
at 1600 h in December in the eastern shadow of the Sierra Nevada Mountains. 


Phaneta lapidana (Walsingham) 
(Figs. 3, 4) 


Semasia lapidana, Walsingham 1879:58, pl. 73, fig. 8 (imago) 

Thiodia lapidana, Fernald [1903]:462; Heinrich 1923: 50; McDunnough 1939:44. 
Eucosma lapidana, Barnes & McDunnough 1917:no. 7085. 

Thiodia lepidana, Heinrich 1929:2, fig. 7 (male genitalia) [missp.]. 

Phaneta lapidana, Powell 1983:33. 


The description of Semasia lapidana was based on one male and two females collected 
at Crooked River near Klamath Lake, Oregon on September 22, 1871. A lectotype for Se- 
masia lapidana Walsingham is hereby designated (Fig. 3): male, “Crooked R. (nr. Fort 
Klamath) Jackson Co., Oregon, 21—23 IX 1871, Wlsm.” BM genitalia slide 11598 (Fig. 4). 
The lectotype is deposited in BMNH. According to Obraztsov’s notes on the envelope 
containing the photograph of the female cotype of S. lapidana, the detached abdomen ac- 
companying this specimen is that of a male. Obraztsov's photograph of this male genitalia 
reveals that it belongs to a specimen of Epinotia columbia (Kearfott), which has been 
treated historically as a junior synonym of the Palearctic species, E. crenana (Hiibner). 
The latter was also collected by Walsingham at Crooked River and misidentified as a 
smaller form of Epinotia (sensu Proteopteryx) emarginana (Walsingham) (1879: pg. 69). 
We examined the following specimen: British Columbia: Chilcotin, 15-IX-1925, George V. 
Copley. 


VOLUME 51, NUMBER 2 121 


Fics. 1-4. Lectotypes of Phaneta species described by Walsingham from Jackson 
County, Oregon. 1, P. sublapidana, male imago; 2, P. sublapidana, male genitalia, B.M. 
slide 11599; 3, P. lapidana, male imago; 4, P. lapidana, male genitalia, B.M. slide 11598. 


Phaneta kokana (Kearfott) 
(Fig. 5) 


Eucosma kokana, Kearfott 1907:29. 

Eucosma chortaea, Meyrick 1912:35 [invalid repl. name]. 

Hystricophora kokana, Heinrich 1923:259 (lectotype designation). 

Thiodia sororiana, Heinrich 1923:263, fig. 421 (male genitalia); McDunnough 1939:44 (as 
subspecies of Thiodia kokana). 

Thiodia kokana, Heinrich 1924:387; McDunnough 1939:44. 

Phaneta kokana, Powell 1983:33; Godfrey et al. 1987:35. 


Eucosma kokana was based on a female from Cincinnati, Ohio and a male from Scran- 
ton, Pennsylvania. Heinrich (1923) designated the female as lectotype and placed the spe- 
cies provisionally in Hystricophora. In the appendix to that same paper, Heinrich described 
Thiodia sororiana and figured the male genitalia based on specimens from Aweme, Mani- 
toba, noting that he had seen specimens of the same species from Ontario, Canada in the Fer- 
nald Collection. He stated that the forewing had a dark band bordering the termen, differenti- 
ating it from P. lapidana, and that it most closely resembled Hystricophora kokana (Kearfott), 
based on examination of a female specimen of the latter. Following receipt of additional 
specimens of P. kokana from Cincinnati, Heinrich (1924) synonymized T. sororiana with 
E. kokana and transferred kokana to Thiodia. To confirm the identity of our specimens, a 
photograph of a female was compared with the lectotype of P. kokana in the AMNH. 

In P. kokana the forewing is divided roughly into a dark basal area and pale apical area, 
the latter extending from beyond the middle of the costa across the wing to the tornus and 
outward to the apex. The basal area bears a mixture of light brown scales, gray scales, and 
gray scales with white or black tips. The white tipped scales are concentrated along the 
costa, and the black tipped scales are distributed evenly except towards the inner margin 
where they are more dense. The varying number of brown scales produces an effect rang- 
ing from a light shading to a predominantly brown color. The apical portion of the wing is 
clothed largely with white and light gray scales with white apices; specimens with exten- 
sive suffusing of brown in the basal area have some brown scales in the apical area as well. 


122 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


A row of darker gray scales forms a conspicuous, straight, terminal line. Some individuals 
show three or four dark marks along the outer half of the costa, which appear to be areas 
accentuated by indistinct, diffuse strigulae. Wing length: males 9.2-11.5 mm (n = 18; 
mean = 10.2; s.d. = 0.47), females 8.3—9.7 mm (n = 3; mean = 9.2). 

This species has a wide distribution, but its scarcity in collections suggests that it is ei- 
ther localized or overlooked because of its late flight period. We examined the following 
specimens: Canada. Manitoba: Aweme, 22 Sept 1921, N. Criddle (1d, genitalia slide 
USNM 70001). Ontario: London, (4¢; genitalia slide USNM 70000). United States. Ken- 
tucky: Rowan Co., E side Rt. 1274, 2 mi W of Rt. 519, 8 Nov 1994, L.D. Gibson (12). IIli- 
nois: Putnam Co., 22 Oct 1946, M. O. Glenn (1d), 25 Oct 1946 (14; genitalia slide MOG 
219), 10 Oct 1949 (1d), 16 Oct 1949 (2d: genitalia slide DJW 103), 18 Oct 1949 (2c), 29 
Oct 1950 (2d), 28 Oct 1964 (1d), 24 Oct 1974 (1d). Massachusetts: Hampshire Co., 
Amherst, Goodell, Fernald Coll. (24, 19; 2 genitalia slide DJW 102). Ohio: Adams Co., 1 
mi SE Lynx, 24 Oct 1991, L. D. Gibson (34; genitalia slide LDG 161), 25 Oct 1991, D. J. 
Wright (1d; genitalia slide DJW 48), 4 Nov 1994 (20d, 5°), Athens Co., Ames Twn., 31 Oct 
1990, H. T. Enterline (2d; genitalia slides DJW 49, 50); Hamilton Co., Cincinnati, 3 Oct, 
A. Braun (2, lectotype, genitalia slide C.H.), 7 Nov 1918, A. Braun (1d, 19; d genitalia slide 
USNM 69998, 2 genitalia slide USNM 69999). Pennsylvania. Lackawanna Co., Scranton, 
8 Nov 1905, A. E. Lister (1d, paralectotype). 


Phaneta ambodaidaleia Miller 
(Fig. 6) 


Phaneta ambodaidaleia, Miller 1983:101, figs. 12-14 (imago, female and male genitalia); 
Miller 1987:47 (figs. imago, male and female genitalia). 


Miller (1983) described this species from Kentucky (Oldham Co.), Michigan (Ingham 
Co.), Missouri (Jasper Co.), North Carolina (Carteret Co.), and South Carolina 
(Charleston Co., type locality). The forewing ground color is creamy white, being most ev- 
ident between the costa and radial vein. Between the radial and cubital veins, it has 
brownish ochreous longitudinal streaks extending from the base outward through the cell. 
From the distal edge of the cell to the termen the brownish ochreous streaks are between 
the veins, accenting the creamy white on the veins. In most specimens the area between 
the cubitus and inner margin is suffused with brownish gray. The forewing is overlaid with 
a sprinkling of dark brown dots, accentuated on the apical half of costa and the outer mar- 
gin. The dots on the outer margin occur between veins, creating the impression of an in- 
termittent terminal line. Wing length: males 9—10.5 mm (n = 24; mean = 9.1; s.d. = 0.33); 
females 7.9—8.3 mm (n = 4; mean = 8.1). 

We add the following records: Alabama: Baldwin Co., Bon Secour N.W. Ref., T9S, 
R2E, Sec 24, 18 Jan 1993 (1d). Georgia: Clinch Co., DuPont, 19 Feb 1983 (12). Kentucky: 
Bullitt Co., N side Rt. 480, 6.9 mi E Rt. 61, 30 Mar 1993, L. D. Gibson (64; wing slide 
LDG 1), D. J. Wright (1d). Mississippi: Hancock Co., Stennis Space Center, 27 Jan 1993, 
R. Kergosien (1d); Harrison Co., Long Beach, 6 Feb 1994, R. Kergosien (3d); Lee Co., 
Tombigbee State Park, 10-31 Mar 1993, R. Kergosien (1d); Oktibbeha Co., 6 mi SW 
Starkville, 24 Feb 1985, R. L. Brown (1¢, genitalia slide RLB 1665), 2 Mar 1985 (4¢), 3 
Mar 1985 (1d), 9 Mar 1986 (1d), TI8N,RI4E, Sec 33 SE, 2 Mar 1991 (3d), 4 Mar 1991 
(1d), Mississippi State University North Farm, 6 Mar 1991 (1°), TION,RI5E. Sec 16 
[Black Belt Prairie], 5 Mar 1991, D. M. Pollock (1d). Ohio: Adams Co., Lynx Preserve, 20 
Mar 1991, D. J. Wright (4d; genitalia slides DJW 47, 74), 1 mi SE Lynx, 20 Mar 1991, L. 
D. Gibson (4¢; genitalia slide LDG 162), 7 Apr 1992 (3d), 26 Mar 1993, L. D. Gibson (1¢), 
D. J. Wright (43), 8 Apr 1993, D. J. Wright (4¢,12). South Carolina, Charleston Co., Mc- 
Clellanville, 20 Mar 1968, R. W. Hodges (22; genitalia slides DJW 82, 102). 


Phaneta canusana Wright, new species 
(Figs. 7-9) 


Description. Head: Scales on vertex, upper frons, and labial palpi brownish gray, shad- 
ing toward white at their bases, with distinctly white tips. Antennae finely pubescent ven- 


VOLUME 51, NUMBER 2 1278} 


Fics. 5-7. Males of Phaneta species from Adams County, Ohio. 5, P. kokana; 6, P. am- 
bodaidaleia; 7, P. canusana, holotype. 


trally, covered dorsolaterally with narrowly white-tipped, brownish gray scales. Thorax: 
Mesonotum and tegulae concolorous with head. Forewing (Fig. 7). Wing length: males 
7.1-10 mm (n = 39; mean = 8.6; s.d. = 0.64), females 6.9—7.6 mm (n = 4; mean = 7.3). 
Dorsal vestiture a mixture of gray to brownish gray scales, mostly tipped with white, pro- 
ducing a unicolorous ashy gray appearance; some individuals with darker scales between 
veins and lighter scales on veins producing weakly highlighted veins and striate appear- 
ance; outer margin of wing with thin, distinct, dark gray terminal line at edge of wing 
membrane, accentuated by white bases of scales in basal row of fringe; fringe scales 
brownish gray with white apices. Hindwing: Upper side and fringe scales uniformly light 
brownish gray, with darker scales usually accenting the veins and wing margins. Male gen- 
italia (Fig. 8): Tegumen widened dorsally; uncus reduced to rudimentary setose lobe; socii 
short, slightly flattened, with lateral margin convex, median margin concave; aedeagus 
short, with more than 20 cornuti; juxta with short caulis, anellus not closely surrounding 
aedeagus ventrally; valva with base of sacculus sparsely setose, with large group of dense 
setae on basal medial area, neck sparsely setose on ventral margin, cucullus sub-triangular 
with median surface angled from dorsal edge of valva and overlapping neck ventrally 
(13n). Female genitalia (Fig. 9): Sternum VII densely scaled on anterolateral and postero- 
lateral comers and medial area anterior to ostium, sparsely scaled elsewhere, anteriorly 
rugose; tergum VIII sparsely setose on lateral extensions and posterior half of dorsum, 
scales absent; papillae anales facing laterally, setae sparse on medial and dorsal areas of 
pads, more dense on ventral margins; lamella postvaginalis with lateral margins slightly 
concave, with shallow longitudinal groove medially, with four or five setae in irregular pat- 
tern, microtrichiate throughout; ductus bursae with moderately sclerotized colliculum 
posterior to inception of ductus seminalis; width of smaller signum less than one half 
greatest width of larger signum (2n). 

Types. Holotype 3, “OH: Adams Co., Lynx Prairie Preserve, Station 6, March 17, 1989, 
leg. D. J. Wright.” Type locality at 38°45'40"N 83°24'46"W. The holotype is deposited in 
USNM. Paratypes. Kentucky: Bullitt Co., Co. Rd. 480, 7 mi E of Shepherdsville, 30 Mar 
1993, D. J. Wright (14, 12; 2 genitalia slide DJW 104), N side Rt. 480, 6.9 mi E Rt. 61, 30 
Mar 1993, L. D. Gibson (1d); Rowan Co., E side Rt. 1274, 2 mi W Rt. 519, 13 Mar 1995, 
L. D. Gibson (9d, 22). Mississippi: Oktibbeha Co., TI9N,RI5E, Sec. 16 [also known as 
“Osborn Prairie”; 33°30'41"N 88°44'08"W], 10 Feb 1991, Black Belt Prairie, D. M. Pol- 
lock (5d), 13 Feb 1992, R. L. Brown (5d), 2 Feb 1995, R. L. Brown (1d). Missouri: Benton 
Co., 3 miles NW of Warsaw on State Hwy UU [data are inaccurate as Hwy UU intercepts 
State Highway 7 at 5 miles NW of Warsaw], 28 Feb 1976, J. R. Heitzman (1d; genitalia 
slide D. Hagler 924803), 2 Mar 1976, J. R. Heitzman (2d; genitalia slides D. Hagler 
610801, 729802). Ohio: Adams Co., Lynx Prairie Preserve, 17 Mar 1989, D. J. Wright (14, 
12, 3 genitalia slide LDG 78, 2 genitalia slide W. E. Miller 1112921), 1 mi. S. E. of Lynx, 
20 Mar 1991, L. D. Gibson (4¢, genitalia slide LDG 160), D. J. Wright (1d, genitalia slide 
DJW 76), 2 Mar 1992, D. J. Wright (76; genitalia slides DJW 51, 52, 75, W. E. Miller 


124 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 8. Phaneta canusana, male genitalia. Adams County, Ohio, slide W. E. Miller 
1112922. Scale line 0.5 mm. 


812922, 1112922), 4 Mar 1992, L. D. Gibson (3¢; wing slide LDG 2), 26 Mar 1993, L. D. 
Gibson (1d), D. J. Wright (53), 8 Apr 1993, D. J. Wright (3d). South Carolina: Greenville, 
23 Feb 1982, Richard S. Peigler (1d; genitalia slide R. L. Brown 1565). Paratypes are de- 
posited in collections of DJW, Cincinnati, OH, LDG, Florence, KY, JRH, Independence, 
MO, AMNH, CNC, MEM, USNM, ZM, Copenhagen [R. Peigler donation]. 

Diagnosis. Phaneta sublapidana and P. lapidana are western species with fall flight pe- 
riods. Heinrich (1923) considered them to be closely related, apparently based on Wal- 
singham’s descriptions and an examination of a cotype of Semasia sublapidana. He sepa- 
rated them in his key by Walsingham’s description of the setation of the male antennae, 
stating that it is “strongly pubescent” in suwblapidana and “nearly smooth” in lapidana. He 
later obtained specimens of P. lapidana from British Columbia and commented (1929) 
“lepidana |sic] resembles kokana Kearfott, which may be nothing but an eastern variety.” 
The male specimen of P. sublapidana that we examined has dense setae covering about 
two-thirds the circumference of each flagellomere, the setae being subequal in length with 
the width of the flagellomere. The female of P. sublapidana has sparce setae restricted to 
less than a third of the flagellomere circumference, and the setae are much shorter than 
the flagellomere’s width. The male specimen of P. lapidana that we examined has a narrow 
strip on the ventral surface of each flagellomere covered with setae that are subequal in 
length with the width of the flagellomere. The remainder of the circumference of the fla- 
gellomere is covered with white scales. Although the forewing ground color is similar in 
both species, P. sublapidana differs from P. lapidana and P. canusana in having a contrast- 
ing light apical area beyond the discal cell in which light gray scales are intermixed with 
grayish orange and light brown scales. Phaneta sublapidana differs from the other species 
treated here in having male genitalia with a deep and wide ventral emargination of the val- 
val neck (Fig.2). Phaneta lapidana is similar to P. canusana in both forewing color and 
male genitalia. 

Phaneta kokana, P. canusana and P. ambodaidaleia are eastern species that are sym- 


VOLUME 51, NUMBER 2 125 


Fic. 9. Phaneta canusana, female genitalia. Bullitt County, Kentucky, slide D. J. 
Wright 104. Scale line 1.0 mm. 


126 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


patric in at least part of their ranges. They have been collected at a single site in Adams 
Co., Ohio. Their genitalia are similar. In males of P. ambodaidaleia, the basal portion of 
the cucullus that overlaps the valval neck has a convex margin. This margin is straight from 
the ventral comer of the cucullus to the neck in the other two species. Wing color and 
flight period provide reliable means of differentiating the three species. The ground color 
of the forewing in P. ambodaidaleia is creamy white, and that of P. kokana and P. canusana 
is gray. In P. kokana the ground color is suffused with light brown scales in the basal area 
and white or white tipped scales in the apical third of the wing, thereby distinguishing its 
appearance from that of P. canusana, which is unicolorous gray. Mean forewing lengths of 
males of the three species differ significantly, with P. canusana being the smallest and P 
kokana the largest. The flight period of P. kokana varies from late September to early No- 
vember, depending on latitude, whereas P. ambodaidaleia and P. canusana fly in the 
spring. Earliest capture dates for P. ambodaidaleia range from January on the Gulf Coast 
in Alabama and Mississippi to February 24 in Oktibbeha County, Mississippi and March 
20 in Adams County, Ohio. In Ohio the flight period of P. canusana begins one to two 
weeks earlier than that of PR. ambodaidaleia and partially overlaps the latter in late March 
and eal April. A similar staggering of flight periods occurs in Mississippi, although no 
overlap has yet been recorded. 

Habitat. Phaneta canusana appears to be Aesedl with a habitat supporting prairie 
vegetation, based on extensive collecting in various physiographic regions in Ohio, Ken- 
tucky, Missouri, and Mississippi. The Lynx Prairie Preserve and adjacent collection site 
one mile SE of Lynx are among an extensive group of small openings in the otherwise 
forested ridge tops and valleys of the unglaciated portion of southern Ohio. The Kentucky 
sites in Rowan and Bullitt Counties are similar, but do not possess the plant diversity of 
the Ohio localities. The Missouri specimens come from the vicinity of the Osage Plains, 
which contains many of the state’s native prairies, and the single South Carolina specimen 
was collected at a porch light in a developed suburb of Greenville. The latter locality was 
originally wooded, and in 1982 the nearest open habitat supporting prairie plants was 
about three miles distant from the collection site. 

The preference for a prairie habitat is supported by collections in the Black Belt phys- 
iographic region of Mississippi, an area discretely defined by an underlying layer of Selma 
Chalk that extends in a crescent shape from northern Mississippi to southeastern Alabama. 
Nine prairie remnants in the Black Belt of Mississippi have been sampled with blacklights 
on 93 nights since 1990; of these, three remnants were sampled on 16 nights in January, 
February, and March. Phaneta canusana was collected at the Osborn prairie remnant on 
three of the four nights during February but not in January or March. Neither P. canusana 
nor P. kokana was collected at this site during eight nights in September, eight nights in 
October, and four nights in November. Other types of habitats in the Lower Coastal 
Plains, Jackson Prairie, Alluvial Plain, Loess Hills, and Flatwoods physiographic regions 
(as mapped in Testa and Lago, 1994) have been extensively sampled over many years dur- 
ing January—March without producing records of the new species. Larval hosts have not 
been recorded for any of the five species discussed in this paper, but they probably are 
Asteraceae, as is the case with other species in the genus. 


ACKNOWLEDGMENTS 


The first author would like to thank W. E. Miller and J. A. Powell for advice and en- 
couragement. The first Ohio record of P. canusana was produced during a survey, sup- 
ported by the Ohio Chapter of The Nature Conservancy, of the Lepidoptera of the Lynx 
Prairie Preserves, a 53 acre tract maintained by The Cincinnati Museum of Natural His- 
tory. Additional specimens of all three species were later recorded at an adjacent site be- 
longing to P. Knoop, who graciously granted permission to collect on his property. Speci- 
mens from Mississippi and Alabama were collected with support from NSF grants 
BSR-9024810 and DEB-9200856 (to R. L. Brown). Collecting permission and support 
from Jerome Carroll and Joe Hardy, Managers of Bon Secour National Wildlife Refuge, 
and Robert Griffin, of the Mississippi Department of Wildlife, Fisheries and Parks is ap- 
preciated. Drawings of genitalia were made by Joe MacGown. Thanks also to B. Landry 


VOLUME 51, NUMBER 2 are 


and J.-F. Landry for providing data from the CNC, R. W. Hodges of USNM and K. 
Methven of INHS for the loan of specimens, and Sidney MacDaniel and the Mississippi 
Natural Heritage Program for habitat information. The contribution by R. L. Brown was 
approved for publication as Journal Article No. J-8846 of the Mississippi Agricultural and 
Forestry Experiment Station, Mississippi State University, with support from State Project 
MIS-6537. 


LITERATURE CITED 


BARNES, W. & J. MCDUNNOUGH. 1917. Checklist of the Lepidoptera of Boreal America. 
Herald Press, Decatur, Illinois. 392 pp. 

FERNALD, C. H. [1903]. In H. G. Dyar, A list of North American Lepidoptera. U.S.N.M. 
Bull. 52:1—723. 

GODFREY, G. L., E. D. CAsHaTT & M. O GLENN. 1987. Microlepidoptera from the Sandy 
Creek and Illinois River Region: an annotated checklist of the suborders Dacnonypha, 
Monotrysia, and Ditrysia (in part) (Insects). Il. Nat. Hist. Surv. Spec. Publ. 7:1—44. 

HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucosminae 
of the family Olethreutidae. U. S. Nat. Mus. Bulletin 123:1—298. 

. 1924. North American Eucosminae, notes and new species (Lepidoptera). J. 

Washington Acad. Sci. 14:385—393. 

. 1929. Notes on some North American moths of the subfamily Eucosminae. Proc. 
Wes Nat. Mus. 75:1—23. 

KEARFOTT, W. D. 1907. New North American Tortricidae. Trans. Am. Entomol. Soc. 
5o:1—9s: 

McDuUNNOUGH, J. 1939. Check List of the Lepidoptera of Canada and the United States 
of America. Part II. Microlepidoptera. Mem. So. Cal. Acad. Sci. 2(1):3-171. 

Meyrick, E. 1912. On some impossible specific names in micro-lepidoptera. Entomol. 
Mon. Mag,, Ser. 2, 48:32—36. 

MILLER, W. E. 1983. Genus Phaneta: new synonymies and a new species (Lepidoptera: 
Tortricidae). Ann. Entomol. Soc. Am. 76:98—103. 

. 1987. Guide to the olethreutine moths of midland North America (Tortricidae). 
U.S.D.A. For. Serv. Agric. Handbook 660:1—104. 

OBRAZTSOY, N. O. 1952. Thiodia Hb. as not a North American genus (Lepidoptera, Tor- 
tricidae). Entomol. News 63:145-—149. 

POWELL, J. A. 1983. Tortricoidea, pp. 31-42. In R. W. Hodges et al. (Eds.), Check list of 
the Lepidoptera of America north of Mexico. E. W. Classey & Wedge Entomol. Res. 
Foundation, London. 

Testa, S. & P. K. Laco. 1994. The aquatic Hydrophilidae (Coleoptera) of Mississippi. 
Miss. Agric. For. Exp. Sta. Tech. Bull. 193:1—71. 

WALSINGHAM, LORD. 1879. Illustrations of typical specimens of Lepidoptera Heterocera 
in the collection of the British Museum. Part IV. North American Tortricidae. 84 pp. 
+ 17 pls. Dept. Zoology, British Museum, London. 


Received for publication 5 October 1995; revised and accepted 1 April 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 128-134 


THREE ADDITIONAL BACTRA IN CALIFORNIA, 
ONE NATIVE BUT OVERLOOKED, ONE PROBABLY 
INTRODUCED, ONE NEW SPECIES (TORTRICIDAE) 


JERRY A. POWELL 


Essig Museum of Entomology, University of California, 
Berkeley, California 94720, USA 


ABSTRACT. = Bactra maioriana Heinrich, which was not recorded west of Utah pre- 
viously, B. priapeia Heinrich, a Gulf Coast species believed to have been introduced into 
coastal southern California, and B. miwok, new species (TL: China Camp, Marin Co.) are 
recorded. Diagnostic features for these and the two previously recorded species, B. 
verutana Zeller and B. furfurana (Haworth) are illustrated. Adults of B. miwok appear to 
be diurnal; early instar larvae mine leaves of Cyperus and presumably Carex. 


Additional key words: Olethreutinae, range expansion. 


Members of the genus Bactra Stephens (Tortricidae: Olethreutinae, 
Bactrini) are relatively narrow winged, mainly tan tortricids that resem- 
ble crambids when at rest. Larvae of several species in the Palaearctic, 
Nearctic, Hawaii, and India, feed in stems of Carex, Cyperus, Scirpus 
and other sedges (Cyperaceae), Juncus (rushes, Juncaceae), and Typha 
(cat-tails, Typhaceae) (Emmet 1979, Frick & Garcia 1975, Heinrich 
1926, Fletcher 1932, Zimmerman 1978). There are more than 50 de- 
scribed species of Bactra, distributed primarily in pantropical and sub- 
tropical regions. Most of the species are both variable in forewing pat- 
tern and similar to one another, so that study of the genital structures is 
necessary to differentiate them. Diakonoff (1956, 1962, 1963, 1964, 1973) 
has illustrated most of the species with excellent, detailed drawings. 

Parallel variation in forewing pattern was categorized and illustrated 
with photographs by Diakonoff (1962). The forewings show a complex 
pattern of pale and dark transverse markings (‘fasciate type’, considered 
by Diakonoff to be primitive and retained only in B. furfurana) or 
within a species often they are unicolorous tan or brown or may feature 
variable maculation (‘maculate type’) and/or a broad, dark dorsal half or 
median band from base to apex (‘vittate type’). Most species display at 
least the unicolorous and vittate types. 

Six species of Bactra occur in America north of Mexico, of which two 
have been recorded in California (Heinrich 1926, Diakonoff 1964): 

Bactra verutana Zeller (TL: Dallas, TX) occurs throughout much of 
North America and is recorded in Puerto Rico, Paraguay, and South 
Africa (Diakonoff 1964). In California it is widespread at lower eleva- 
tions, including coastal areas, Central Valley, and deserts (CAS, CDFA, 
LACM, UCB, UCD; see acknowledgements). The California race was 
designated as chrysea Heinrich (1926), but as pointed out by Diakonoff 
(1964), there is no justification in applying a subspecies name to any 


VOLUME 51, NUMBER 2 129 


subset of these variable populations. B. verutana often is numerous at 
lights and presumably is multivoltine (e.g., there are records for every 
month from April to December in the San Francisco East Bay area). 
Most collections have been made in July, August and September. The 
adults are small (male forewing length 5.6—8.4 mm, female 6.8—9.5 
mm) and usually have a more or less unicolorous tan forewing or with 
reduced blackish maculation; a few exhibit the ‘vittate type’ pattern with a 
longitudinal dark streak. The genitalia of both sexes are diagnostic (Figs. 
1, 6). The larvae feed in sedges, especially Cyperus, including weedy spe- 
cies, and the biology has been extensively studied in relation to possible 
biological control of nutsedges (Poinar 1964, Frick & Garcia 1975). 

Bactra furfurana (Haworth) (TL: England) is a Holarctic species that 
also occurs widely in California but is more localized and less often en- 
countered. Most of the records are more northern (North Coast, inland 
Mendocino Co., Sacramento Valley), although B. furfurana has been 
taken at La Jolla, San Diego Co. and Bishop, Inyo Co. (UCB). It appears 
to be absent from Lower Sonoran zones in the San Joaquin Valley and 
desert areas. The voltinism is not documented in California; collection 
records range from late April (Hopland Field Sta., Mendocino Co.) to 
mid July (Bishop), but there has not been seasonal monitoring at one lo- 
cality. I have not seen B. verutana and B. furfurana occurring in close 
sympatry. The forewing, which is dark with variable pale markings (‘fas- 
ciate type’) and has a more acute apex, distinguishes B. furfurana from 
the other California species, as do the genitalia (Figs. 2, 7). 

Three additional species have been confirmed as resident in Califor- 
nia: Bactra maioriana Heinrich, which is a widespread but rarely col- 
lected, presumably native species; B. priapeia Heinrich, likely an intro- 
duced species in coastal southern California; and B. miwok, new species, 
the male of which has been known for 35 years, the female only recently. 


Bactra maioriana Heinrich 
Bactra maioriana, Heinrich 1923, Proc. Entomol. Soc. Washington, 25:105 (TL: VA). 


This species is a widespread, apparently native, Nearctic insect. Diakonoff (1964) pre- 
sented illustrations of the diagnostic genitalia and regarded B. maioriana to be distinct 
from any Old World species. The species is recorded in the Great Lakes region (Miller 
1987) and westward to North Dakota (UCB) and Utah (Heinrich 1926). The larvae are re- 
ported to feed in stems of Scirpus and Typha (Heinrich 1926, Diakonoff 1964). 

The Utah records, paratypes collected at Vineyard in 1914 and 1922, and Bear River 
Bay, Great Salt Lake (Braun 1925), are considered to represent native populations. No 
other western representatives were recognized until we took specimens at the Antioch Na- 
tional Wildlife Refuge, Contra Costa Co., Calif., in July 1990. At this site, both Scirpus cal- 
ifornicus (C. Meyer) and Typha latifolia L. grow along the San Joaquin River. Because this 
is a shipping route, an introduction via cargo ships in recent times seemed to be a plausi- 
ble explanation for the discovery of this disjunct population. During search of collections, 
however, I recovered additional records of B. maioriana from disparate parts of Califor- 
nia. The widespread occurrence suggests that this species is native in the Pacific States but 


130 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-5. Male genitalia of Bactra species. 1, B. vertutana Z., valva, inner aspect 
(JAP prep. 6413, Bowman, Placer Co., Calif.); 2, B. furfurana (Haw.), valva (redrawn from 
Diakonoff 1956, Europe); 3, B. maioriana Heinr., valva (redrawn from Diakonoff 1964, 
paratype); 4, B. priapeia Heinr., valva and a, aedeagus drawn to same scale (redrawn from 
Diakonoff 1964, holotype); 5, B. miwok Powell, ventral aspect, aedeagus removed and 
shown dorsolateral aspect (holotype). [scale = 0.5 mm for Fig. 5]. 


has been overlooked, perhaps in part owing to its superficial similarity to the more abun- 
dant B. verutana. Individuals of maioriana tend to be larger (male 8.2 mm forewing 
length, female 8.5—10.5 mm in California examples) and darker colored, with a greater 
frequency of vittate type, but there is overlap in forewing phenotype. The two are sym- 
patric at Antioch and Bundy Canyon, Riverside Co. Available records (Diakonoff 1964, 


VOLUME 51, NUMBER 2 Sy 


Miller 1987, present data) suggest a univoltine cycle, adults flying in late spring to early 
summer. 

The male genitalia are similar to those of B. verutana and B. miwok (see diagnosis of 
latter, Figs. 1,3,5), while the female has distinctive, heavily sclerotized, rugose patches lat- 
erally in the VIII-IX intersegmental membrane (Fig. 8). 

California data: Contra Costa Co.: Antioch National Wildlife Refuge, 16, 12 VII-10-90, 
at blacklight (Y.-F. Hsu & J. Powell, UCB) [JAP genit. preps. 6371, 6372]. Monterey Co.: 
Castroville, 12 VI-2-61 (W. H. Lange, UCD) [JAP prep. 6420]. Riverside Co.: Bundy Cyn. 
1660', 9 mi. S Perris, 12 VII-20-76 (R. J. Ford, LACM) [JAP prep. 6405]. San Luis Obispo 
Co.: La Panza Campground, 12 mi. NE Pozo, 1° V-2-62 [at lights] (J. Powell, UCB) [JAP 
prep. 6449]. 


Bactra priapeia Heinrich 
Bactra priapeia, Heinrich 1923, Proc. Entomol. Soc. Washington 25:105 (TL: LA). 


This species has been recorded from Florida, Louisiana, the Gulf Coast of Texas, 
British Honduras, and Panama (Heinrich 1926, Diakonoff 1964). The adults are larger 
than any other North American Bactra (male 9.3 mm forewing length, female 10.1—-10.9 
in California examples), and the genitalia are markedly distinct in both sexes (Figs. 4, 9). 
Forewings of California specimens are pale, rusty, or dark tan without distinct markings, 
but maculate and vittate examples are reported by Diakonoff. The larval hostplant has not 
been recorded. 

During a light trapping survey by the San Diego County Department of Public Health 
in 1959, R. A. Mackie recovered one female of B. priapeia at Mission Bay, an area subse- 
quently converted from marshland to parks. Although I identified the species in 1960, it 
was not until 1987 that another record was obtained to confirm the residency of this spe- 
cies in California. Because the previous collection records are from areas adjacent to the 
Gulf of Mexico, it seems likely that B. priapeia is introduced in coastal southern California. 

California data: Orange Co.: Aliso Cr., 0.5 km E Highway 1, nr. South Laguna, 1d, 2° 
X-7/8-87, at lights (J. Powell, UCB) [JAP genit. preps. 6445, 6446]. San Diego Co.: Mis- 
sion Bay, 1° I[X-30-59, light trap (R. A. Mackie, UCB) [JAP prep. 501]. 


Bactra miwok Powell, new species 


Description. A small, almost uniformly dark brown species with only indistinct ochre- 
ous mottling on the forewings of some individuals. Male: forewing (FW) length 5.9—7.9 
mm (30n); length 3.2—3.4 x width; termen strongly angled, nearly straight, very slightly 
convex, apex produced. Color of head and palpi concolorous with FW, in fresh specimens 
variable, dark brown tinged with dark ochreous-tan to dark rust-brown; pale examples 
faintly irrorate with pale ochreous, not defining a visible FW pattern. No ‘maculate’ or ‘vit- 
tate’ polymorphism known. FW underside gray, fringe pale rust-ochreous. Hindwing en- 
tirely dark gray, fringe pale gray, underside the same. Abdomen scaling dark shining gray, 
venter slightly paler gray. Genitalia (Fig. 5, drawn from holotype, JAP prep. no. 6390; 5n), 
similar to B. maioriana and verutana, with the major spurs of the cucullus slender, nu- 
merous (ca. 30), and broadly distributed along distal margin and posteriorly; major spurs 
of the valva basally fewer (10—12), also slender; saccular margin broadly expanded later- 
ally. Female: FW length 6.3—7.0 mm (2n), length to width ratio within range of male. Phe- 
notype similar to male, color probably comparably variable. Genitalia (Fig. 10, drawn from 
JAP prep. 7157; 2n), similar to B. verutana, but the ring around IX and lateral patches on 
the intersegmental membrane of that species only weakly sclerotized, lateral lobes more 
widely separated and displaced anteriorly from sterigma. 

Types. Holotype 3: CALIF: China Camp, Marin Co., April 18, 1959 (J. Powell) [JAP 
genit. prep. 6390]. Allotype °: CALIF: Miwok Meadow, China Camp St. Park, Marin Co., 
March 30, 1995 (J. Powell) [JAP prep. 7089]; both in UCB. Paratypes (30): CALIF: Marin 
Co.: same data as holotype, 16 [JAP prep. 555]; Miwok Mdw., China Camp St. Park, 2d 
[11-30-94, 124 V-2-94, 9¢, 1° I1I-30-95 (J.Powell) [JAP preps. 7088 ¢, 7157 °]; Ring Mt., 
1d I1-30-85, 1d TV-11-94 (J. Powell). Monterey Co.: Asilomar, 16 V-17-59 (G. I. Stage) 


132 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 6-10. Female genitalia of Bactra species: VIII-IX segments, sterigma, scleroti- 
zation of intersegmental membrane, and basal portion of ductus bursae, ventral aspect. 6, 
B. verutana (JAP prep. 6950, Coachella Vy., Calif.); 7, B. furfurana (Haw:) (JAP prep. 
7194, Smartville, Calif.); 8, B. maioriana Heinr. (redrawn from Diakonoff 1964, paratype); 
9, B. priapeia Heinr. (redrawn from Diakonoff 1964, Texas); 10, B. miwok Powell (JAP 
prep. 7157, China Camp). [scale = 0.5 mm for Fig. 10]. 


VOLUME 51, NUMBER 2 133 


[JAP prep. 7158]. Santa Clara Co.: 3 mi W New Almaden, 146 IV-18-69 (P. A. Opler) [JAP 
prep. 7162]. Sonoma Co.: Bodega, 14 V-3-36 (E. C. Johnston, USNM). Paratypes will be 
deposited in CDFA, CNC, LACM, Mississippi State U., U. Minnesota, USNM. 

Diagnosis. The male genitalia of B. miwok are similar to B. verutana and B. maiori- 
ana, but the new species has more major spines of the cucullus (ca. 30), which are more 
broadly distributed distally and extended to the anterior margin; in verutana and maiori- 
ana these spines are less numerous (10—12), much stouter and arranged in two separated 
lines in verutana (Fig. 1), restricted narrowly to the distal margin and not reaching the 
posterior margin in maioriana (Fig. 3). The major spines of the basal part of the valva are 
quite short in verutana, varying from 6—8 to 15-20 among individuals, while they are 
longer and slender in maioriana and miwok, fewer in the latter (12—14) than in maioriana 
(22-24). The distal margin of the sacculus is more broadly expanded laterally in miwok 
than in the other two species, which have attenuated marginal extensions. In female gen- 
itla and associatred structures, the new species most closely resembles B. verutana (Fig. 
6), with the sclerotized areas lateral to the sterigma reduced compared to that species, the 
lateral cup-like structures more widely separated and displaced anteriorly to the sterigma 
(Fig.10). The other three species discussed differ greatly in these structures (Figs. 7—9). 

Biological Notes. The new species apparently is univoltine, available records of adults 
spanning late March to mid May. There has not been thorough survey to preclude a sum- 
mer generation, but I made visits to Ring Mountain in late May, July, and September, and 
Miwok Meadow in late August. The vegetation was very dry and no Bactra were recov- 
ered from either site. 

Lack of light attraction records suggests that this species is diurnal. The moths were 
netted during midday and early afternoon, and a few were observed to fly in direct sun- 
shine without observer disturbance. Efforts to observe adults during crepuscular and mat- 
inal hours produced negative results. I visited the Miwok Meadow colony site on 10 April 
1994 between 1810-1910 hr PST, just before and after sundown, and on 24 May 1994 at 
0530—0615 hr PST. Observations of caged adults (5d, 12) were inconclusive as to the nor- 
mal diel periodicity. Adults remained ‘alert’ with the antennae erect during daylight hours 
and occasionally actively ran towards the light. It appeared they became more active in 
late afternoon sunshine and at dusk, but several moved after dark. Neither mating nor 
oviposition was observed, but the female deposited about 100 eggs during a 5-day period. 

At Miwok Meadow adults occurred in a wet seepage meadow above tidal marshes along 
San Francisco Bay, in association with three species of Carex (C. multicostata Mackenzie, 
C. praegracilis Boott, and C. tumulicola Mackenzie) and Juncus Ppatens E. Meyer, while 
at Ring Mountain they were taken in a narrow seepage gully on a steep slope in serpen- 
tine grassland, with what appeared to be one of the same and a fourth species of Carex, 
and both Cyperus and Juncus. However, no definite association with any one of the plants 
was observed, and pupal shells were not found. None of these plants has a sufficiently ro- 
bust stem to support later instar larvae, particularly through winter, and I assume that late 
instar larvae feed at the base of the plant. 

Early instar larval activity of Bactra miwok in the laboratory suggests that the larval bi- 
ology is similar to that described for B. verutana by Poinar (1964). Eggs of B. miwok were 
deposited in variable clumps of usually 4 to 8 eggs (sometimes singly), usually regularly 
overlapped in pairs so that groups of 4, 6, or 8 were formed. The female, caged with a bou- 
quet of two species of Carex from the field site, selected the semitranslucent, bract-like 
blades that subtend the green foliage as oviposition sites. No eggs were laid on inflores- 
cences nor on the glass sides of the container. Incubation was rapid, requiring 6—7 days 
until hatching at 17—21° C. Poinar reported 3—4 days in B. verutana at 22° C. Provided 
with foliage of Cyperus eragrostis Lam., a native plant that behaves like a weed and has 
much larger leaves than the China Camp Carex species, the first instar larvae tunneled 
downward from cut leaf ends, in straight mines between the leaf veins. The frass-packed 
mines were visible within 2 days of hatching and reached 4—6 cm in length by day 4; by 
day 7 the leaves were drying and had folded along the midrib, and 2nd instar larvae made 
holes to the surface and fed between the folded leaf blade or entered new mines. Accord- 
ing to Poinar (1964), mining in B. verutana extends down to the basal meristematic por- 
tion of the stem, where larvae complete development in 8—11 days in the lab at 24° C. 


134 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Larvae of B. miwok were not maintained on entire plants that would have enabled com- 


pletion of feeding. 


ACKNOWLEDGMENTS 


Cooperation by Richard A. Coleman, San Francisco Bay Refuge Complex, U. S. Fish 
and Wildlife Services, Newark; Kenneth Leigh, California Department of Parks and 
Recreation, San Rafael; and Larry Serpa, The Nature Conservancy, Tiburon, permitted 
survey at the Antioch National Wildlife Refuge, China Camp State Park, and Ring Moun- 
tain Reserve, respectively. I thank the curators of the following collections, whose cooper- 
ation facilitated my search of unidentified accessions and curated material for Bactra: Cal- 
ifornia Academy of Sciences, San Francisco (CAS); California State Department of Food 
& Agriculture, Sacramento (CDFA); Los Angeles County Museum of Natural History, Los 
Angeles (LACM); San Diego Museum of Natural History, San Diego (SDNH); University 
of California, Davis (UCD); Essig Museum, University of California, Berkeley (UCB). 
R. A. Mackie, now of Concord, CA, provided specimens from an extensive light trapping 
program with the San Diego County Department of Public Health in 1959, which are de- 
posited in UCB. Barbara Ertter and Dan Norris, University Herbarium, UC Berkeley, 
provided plant identifications. 


LITERATURE CITED 


BRAUN, A. F. 1925. Microlepidoptera of northern Utah. Trans. Am. Entomol. Soc. 51:198. 

DIAKONOFF, A. 1956. Records and descriptions of Microlepidoptera. Zool. Verhandl., Lei- 
den, 29:1—60. 

. 1962. Preliminary survey of the Palaearctic species of the subgenus Bactra 

Stephens (Bactra, Tortricidae, Lepidoptera). Zool. Verhandl., Leiden 59:1—48. 

. 1963. African species of the genus Bactra Stephens (Lepidoptera, Tortricidae). 

Tijd. v. Ent. 106:285—357; figs. il=78}, 

. 1964. Further records and descriptions of the species of Bactra Stephens (Lepi- 

doptera, Tortricidae). Zool. Verhandl., Leiden 70:1—81. 

. 1973. The south Asiatic Olethreutini (Lepidoptera, Tortricidae). Rijksmus. van 
Natuurlijke Historie. Zool. Monogr. 1. 699 pp. 

EMMET, A. M. (ED.) 1979. A field guide to the smaller British Lepidoptera. Brit Entomol. 
Nat. Hist. Soc., London. 271 pp. 

FLETCHER, T. B. 1932. Life histories of Indian Microlepidoptera. Alucitidae (Pterophori- 
dae), Tortricina and Gelechiadae. Imp. Council Indian Agric. Res., Sci. Monogr., No. 
De ST 

FRICK, KE & GARCIA, C., JR. 1975. Bactra verutana as a biological control agent for 
Purple Nutsedge. Ann. Entomol. Soc. Amer. 68:7—14. 

HEINRICH, C. 1926. A revision of the North American moths of the subfamilies Laspeyre- 
siinae and Olethreutinae. U. S. Natl. Mus. Bull. 132. 216 pp. 

MILLER, W. E. 1987. Guide to the olethreutine moths of midland North America (Tor- 
tricidae). U. S. D. A. Forest Serv., Agric. Handb. 660. 104 pp. 

POINAR, G.O., JR. 1964. Studies on nutgrass insects in southern California and their effec- 
tiveness as biological control agents. J. Econ. Entomol. 57:379—383. 

ZIMMERMAN, E. C. 1978. Insects of Hawaii. Vol. 9. Microlepidoptera, part 1. Univ. Hawaii 
Press, Honolulu. 881 pp. 


Received for publication 13 December 1995; revised and accepted 13 May 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 135-138 


A NEW CHAETAGLAEA FROM THE SOUTHEASTERN 
UNITED STATES (NOCTUIDAE: CUCULLITINAE) 


VERNON ANTOINE BROU, JR. 
74320 Jack Loyd Road, Abita Springs, Louisiana 70420, USA 


ABSTRACT. Chaetaglaea fergusoni, new species, is described and illustrated, and 
compared to other members of the genus. Chaetaglaea tremula and Chaetaglaea sericea 
are reported as new from Louisiana. 


Additional key words: Mississippi, South Carolina, voltinism, winter moths. 


In 1943, Franclemont described the genus Chaetaglaea, listing three 
species: C. cerata Franclemont, C. tremula (Harvey), and C. sericea 
(Morrison). Hodges (1983) retained the same treatment for the genus. 
Both Franclemont (1943) and Forbes (1954) stated that only C. tremula 
ranged south to the Gulf coastal states, and Covell (1984) indicated that 
both C. tremula and C. sericea occurred in the Gulf coastal states from 
Florida to Mississippi. The genus Chaetaglaea has not been reported 
previously from Louisiana by noctuid workers (see von Reizenstein 
1863, Jones 1918, Folsom 1936, Glick 1939, Harrison 1946, Merkl and 
Pfrimmer 1955, Pfrimmer 1957, Chapin & Callahan 1965). The purpose 
of this paper is to describe a new Chaetaglaea from Louisiana, and to 
provide new distributional and phenological data for the genus from the 
southern United States. 


Chaetaglaea fergusoni Brou, new species 
(Figs. 1, 2) 


Description. Male (Fig. 1A): Mean forewing length 19.6 mm (range 18.9—20.4 mm, 
n=13). Head: color medium or reddish brown, with orange hue, frontal tuft rounded; palpi 
color contrasting reddish brown or orange brown; antennae similar matching color, sim- 
ple, slender, acuminate. Thorax: frontal area dorsal color similar to head, orange brown 
scales less numerous caudally, color becoming reddish brown; scales form longitudinal 
central ridge; ventral color reddish brown; legs reddish brown to orange brown, usually 
similar to color of palpi. Abdomen: dorsal color rich reddish brown, on aged specimens 
brown to tan, generously peppered with black scales which substantially increases cau- 
dally; ventral color same as dorsal; anal tuft orange brown. Forewing: dorsal ground color 
rich monochrome reddish brown, light dusting of pale whitish scales basad of postmedial 
line, especially concentrated along costal margin near base; same area peppered with nu- 
merous, well dispersed, inconspicuous, black scales; whitish scales form a barely notice- 
able, thin line distally hugging postmedial line; single reniform spot, dark and small, some- 
times nearly distinguishable or nonexistent; basal line usually limited to dark dash or spot 
on costal margin, occasionally extending onto basal area of discal cell; dark antemedial line 
extending from costal edge to a point about one-third width of wing, directed toward anal 
angle, abruptly changing direction perpendicular to inner margin, increasingly obscure, 
often vanishing; similar medial line roughly parallel to antemedial line, curving basally ap- 
proaching inner margin; similar postmedial line beginning at costal edge, roughly parallel 
to medial line, usually vanishing two-thirds distance to inner margin, but occasionally rep- 
resented by short dash near costal margin or distinct dark line intersecting inner margin; 
subterminal line changes from rich reddish brown ground color to area completely and 


136 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Chaetaglaea fergusoni, new species. A, male holotype; B, female allotype. 


delicately burnished with lighter shiny brown scales, creating whitish suffusion to outer 
margin, distinct to nearly indistinguishable; dark zigzag terminal line inwardly accented by 
tiny black dots between each vein, representing adterminal line, sometimes only adtermi- 
nal dots evident; fringe uncontrasting reddish brown; ventral color fuscous, center bor- 
dered by shiny reddish brown along costal margin and broader area along outer margin; 
maculation limited to postmedial line, prominence variable. Hindwing: dorsal color fus- 


A 8B 


Fic. 2. Genitalia of C. fergusoni, Mississippi, Harrison County. A, B, male (aeodeagus 
at left); C, female. 


VOLUME 51, NUMBER 2 EST 


fergusoni 
sericea 
SUOUCUNUGUNUUEOUUNUNUHNOUN UREN ENUUUNUUNUDUUUMERUOUUUUCGMUNENUOUUNHUUUUUMUGUMUGHMEURUMGHUMUCHUUOUUUUNURONNUUOUCRURUMUORRUOUNNNDOMURUGCOONN NHN 


October November December January February 


Fic. 3. Dates of capture for Chaetaglaea taken at Section 24, Township 6, South 
Range 12 East, 6.8 km NE Abita Springs, Louisiana. Vertical tick marks represent five cap- 
tures: for C. fergusoni, n = 20, 1984-1996; for C. tremula, n = 118, 1990-1995; for C. 
sericea, n = 303, 1990—1995. 


cous postmedially, increasingly lighter tan antemedially approaching base and anal area; 
fringe contrasting light brown, bordered inwardly by fine, inconspicuous, sinuous, reddish 
brown line, itself bordered basally by similar lighter contrasting, fine, tan line; still further 
bordered basally by edge of fuscous wing area, sometimes appearing as distinct adtermi- 
nal line, often accented between some veins as tiny black dots; ventral color shiny reddish 
brown, peppered with well dispersed individual black scales; area near inner margin more 
fuscous; continuation of forewing postmedial line is only maculation. Genitalia (Fig. 2A—B, 
n=3): valves bilaterally asymmetrical; right valve simple, elongate, narrow, cuneate, cucul- 
lus truncate; left valve simple, elongate, narrow, cucullus obtuse; aedoeagus lanceolate, 
curving near obtuse apex, single tiny cornutus; large, elongate, spatulate, free lobes of sac- 
culus extending almost to valve ends; uncus slender, acuminate; saccus greatly elongate, 
tapering to truncate end. Female (Fig. 1B): Mean forewing length 19.4 mm (range 
19.0-19.9 mm, n=5). External morphology as described for male. Genitalia (Fig. 2C, 
n=2): oval unisaccate corpus bursae, small appendix bursae, separated distally by heavily 
sclerotized crescent patch. Sclerotized distal half of ductus bursae. 

Types. Holotype 3 (Fig. 1A): USA, Louisiana, St. Tammany Parish, 4.2 miles (6.8 km) 
NE Abita Springs, Section 24 of Township 6, South Range 12 East, 7 December 1988. Al- 
lotype 2 (Fig. 1B): same locality as holotype, 1 January 1992. Paratypes: 13 5 3 °, same lo- 
cality as holotype, 24 November to 24 February 1984-1996; 5 d 7 °, Mississippi, Harrison 
County, 20 December to 10 January 1992-1993; 1 3, Mississippi, Hancock County, 16 
February 1992; 1 2, South Carolina, Charleston County, Wedge Plantation, South Santee 
River, 22 November 1967. Holotype and allotype deposited at the U.S. National Museum 
of Natural History, Washington, D.C. Paratypes deposited at Louisiana State University, 
Baton Rouge, and in the private collections of Rick Kergosien and the author. 

Etymology. I take pleasure in naming this species in honor of Douglas C. Ferguson, 
who appears to have first discovered it 30 years ago. 

Diagnosis and Discussion. Chaetaglaea fergusoni looks most similar to C. tremula. 
The maculation of both species can be nearly identical, although some specimens of C. 
fergusoni tend to have less accentuated markings. C. tremula is highly variable in both 
color and maculation, but despite this, C. fergusoni can always be separated by its less vari- 
able, rich reddish brown coloration. The male genitalia of C. fergusoni also differ from the 
three other species in the genus. Franclemont (1943) and Forbes (1954) illustrated and 
discussed the male genitalia of C. cerata, C. tremula, and C. sericea: the saccus of each ta- 
pers to an acute apex, the aedoeagus of each possesses large cornuti, and none of these 
three species has large free lobes of the sacculus. Chaetaglaea fergusoni appears to be 
rarely encountered; it has been collected at both ultraviolet light traps and fermenting 


138 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


fruit bait. Apparently univoltine, its winter flight period is influenced by temperature and 
other factors. The flight periods of C. fergusoni, C. tremula and C. sericea at one location 
in Louisiana are shown in F ig. oF 


ACKNOWLEDGMENTS 


I thank the following for supplying specimens, records, or aiding in other ways with this 
project: J. B. Chapin, D. C. Ferguson, L. F. Gall, R. Kergosien, D. F. Schweitzer, and J. B. 
Sullivan. 


LITERATURE CITED 


CHAPIN, J. B. & P. S. CALLAHAN. 1967. A list of the Noctuidae (Lepidoptera, Insecta) col- 
lected in the vicinity of Baton Rouge, Louisiana. Proc. La. Acad. Sci. 30:39—48. 

COVELL, C. V., JR. 1984. A field guide to moths of eastern North America. Houghton Miff- 
lin Co., Boston. 469 pp. 

FOLsoM, J. W. 1936. Notes on little-known cotton insects. J. Econ. Entomol. 29:284—285. 

FORBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Noctuidae, Part 
III. Cornell Univ. Agric. Exp. Sta. Mem. 329:1—433. 

FRANCLEMONT, J. G. 1943. Notes on some Cuculliinae (Phalaenidae, Lepidoptera). III. 
On the identity of Glaea pastillicans Morrison and the species of the genus 
Chaetaglaea new genus. Entomol. News 54:92—99. 

HODGES, R. W. (Ed.). 1983. Check list of the Lepidoptera of America north of Mexico. E. 
W. Classey Ltd. and the Wedge Entomol. Res. Foundation, London. 284 pp. 

GLICK, P. A. 1939. The distribution of insects, spiders, and mites in the air. U.S.D.A. Tech. 
Bull. 673. 

HarRISON, P. K. 1946. Insects attacking cole crops in Louisiana. J. Econ. Entomol. 
39:820-—821. 

JonEs, T. H. 1918. Miscellaneous truck-crop insects in Louisiana. U.S.D.A. Tech. Bull. 
703. 

MERKL, M. E. & T. R. PFRIMMER. 1955. Light trap investigations at Stoneville, Miss., and 
Tallulah, La., during 1954. J. Econ. Entomol. 48:740—741. 

PFRIMMER, T. R. 1957. Response of insects to different sources of blacklight. J. Econ. En- 
tomol. 50:801—803. 

VON REIZENSTEIN, L. 1863. Catalogue of the Lepidoptera of New Orleans and its vicinity. 
Issac. T. Hinton, New Orleans. 8 pp. 


Received for publication 20 May 1994; revised and accepted 24 March 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 139-148 


LARVAL HOSTS OF URESIPHITA HUBNER (CRAMBIDAE) 


ROSEMARY LEEN 


United States Department of Agriculture, Forest Service, 
Pacific Southwest Research Station, P. O. Box 236, Volcano, Hawaii 96785, USA 


ABSTRACT. A survey of the literature and museum collections of Uresiphita indi- 
cates larval hosts are primarily quinolizidine-bearing plants in tribes of the Fabaceae. 
Three species, Uresiphita reversalis, U. ornithopteralis and U. polygonalis, were collected 
from seven genera in the Genisteae (Chamaecytisus, Genista, Lupinus, Spartium, Labur- 
num, Ulex and Cytisus) and from three genera in the Sophoreae (Sophora, Pericopsis and 
Bolusanthus). Two species, U. reversalis and U. polygonalis, were collected from three 
genera in the Thermopsidae (Baptisia, Anagyris and Piptanthus) and two, U. reversalis 
and U. ornithopteralis, were collected from two genera in the Bossiaceeae (Hovea and 
Templetonia). A few legume species that are not known to bear quinolizidine alkaloids 
were also reported. In particular, U. reversalis, U. polygonalis, and U. ornithopteralis were 
each collected from Acacia (Mimosaceae) in areas as widely distributed as Australia and 
the United States (California, Texas and Hawaii). This is a consistent anomaly in the over- 
all host-use pattern. Other nonleguminous species have been reported but are probably 
not indicative of hosts upon which development may be completed. 


Additional key words: Pyralidae, Pyraustinae, aposematism, host plant range, 
French broom, quinolizidine alkaloids. 


In 1983, Uresiphita reversalis (Guenée) caused significant damage to 
Genista monspessulana (L.) L. Johnson, also known as French broom, in 
the San Francisco Bay Area. Thus, U. reversalis was thought to be useful as 
a control agent against the introduced weedy brooms in California (Leen 
1992, 1995). Little was known about the biology and host plant range of 
the genus Uresiphita Hiibner so this survey of collections and publica- 
tions was begun to ascertain if a pattern of host use could be detected. 

Although the genus is in need of taxonomic revision, several species 
and subspecies are recognized and accepted as follows. Uresiphita re- 
versalis, the Genista caterpillar, is the only species known to occur in 
North America (Munroe 1976). Uresiphita ornithopteralis Guenée, the 
tree-lucerne moth, is an Australian species (Common 1990). Several sub- 
species are recognized within Uresiphita polygonalis ({Denis and Schif- 
fermiiller]) by Clarke (1971). Uresiphita polygonalis maorialis (Felder 
& Rogenhofer), the kowhai moth, is indigenous to New Zealand; Ure- 
siphita polygonalis virescens (Butler) is considered indigenous to 
Hawaii but may be introduced (Zimmerman 1958); and Uresiphita 
polygonalis ochrocrossa Clarke is indigenous to Rapa Island (Clarke 
1971). Palm (1986) lists Uresiphita limbalis as a synonym of U. poly- 
gondlis. This paper presents a collation of information available on geo- 
graphical distribution and hosts of these three species of Uresiphita. 


MATERIALS AND METHODS 
Information on the distribution and collections of Uresiphita was ob- 
tained from publications and museum collections. A list of these sources 
appears in Tables 1 and 2. The primary source of information on the dis- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


140 


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JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


142 


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VOLUME 51, NUMBER 2 143 


tribution of species other than U. reversalis was the Natural History 
Museum, London, U.K. Very few host records were associated with 
those specimens, so the majority of host information for all Uresiphita 
species was obtained from the literature and correspondence or visits to 
museums and collections within the United States. Plant species’ names 
are reported as they are currently accepted rather than exactly as re- 
ported on the records. Scientific names, in lieu of common names, are 
reported if no other species or genus could be accorded the common 
names of the associated collection record. 


RESULTS 


Distribution. The genus Uresiphita has been collected from all ma- 
jor continents occurring between 50° north and 50° south latitude. Col- 
lection sites in the northern hemisphere extend into parts of Canada 
(Nova Scotia), the southern part of the United Kingdom and into parts 
of Germany, Poland and the former USSR. Collection sites in the south- 
ern hemisphere extend to New Zealand, South Africa, and the Amazo- 
nian region of Brazil. Collections have also been made from parts of 
western China and several island locations, including Fiji, Norfolk Is- 
land, Rapa Island, the Hawaiian Islands, Madeira, the Canary Islands, 
the Bahamas and San Domingo. Munroe (1976) reported that Ure- 
siphita is found in the Marquesas, although Clarke (1986) made no 
mention of this genus in his volume on the Pyralidae and Microlepi- 
doptera of the Marquesas Archipelago. Munroe (pers. comm.) states 
this was an error on his part. 

Hostplant relationships. Publications and collections of Uresiphita 
indicate all use leguminous species from tribes that are known to contain 
quinolizidine alkaloids (Table 1). These tribes are all within the Fabaceae 
and include the Genisteae, Thermopsidae, Sophoreae and Bossiaceeae. 
Three species, U. reversalis, U. ornithopteralis, and U. polygonalis, were 
recorded from seven genera in the Genisteae (Chamaecytisus, Genista, 
Lupinus, Spartium, Laburnum, Ulex and Cytisus) and from three genera 
in the Sophoreae (Sophora, Pericopsis and Bolusanthus). Two species, U. 
reversalis and U. polygonalis, were recorded from three genera in the 
Thermopsidae (Baptisia, Anagyris and Piptanthus) and two, U. reversalis 
and U. ornithopteralis, were recorded from two genera in the Bossi- 
aceeae (Hovea and Templetonia). Other reported host tribes within the 
Fabaceae include the Phaseoleae (Phaseolus), Trifolieae (Trifolium) and 
the Carmichaeliae (Carmichaelia) (Table 1). The latter fabaceous tribes 
are not known to contain quinolizidine alkaloids. 

Native host plants of U. reversalis include Lupinus, Baptisia and 
Sophora and introduced hosts include Genista and Spartium (Table 1). 
Cytisus scoparius (L.) Link is an introduced plant that is also reported 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


144 


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JEN SNA PEN “6 ‘OL6T POrUNY “g ‘8L6T Avan “1 ‘OZET PreUOAT “9 ‘Q96T UPISYD ‘S ‘LOGIT HeSBO1N “F ‘EBT SEqIOY “E “VSN “OBy pure 
poog ydeq FyeD ‘SZ ‘eg6l Houry ‘T :soommog ‘dds nyydisarp jo syurzd ysoy snoutumsey, uou Jo suoMseTJoo pue sp1ooce1 poysyqng °% ATAVL 


VOLUME 51, NUMBER 2 145 


as a host of U. reversalis but these are not credible records (Leen 1992, 
1997). Reported hosts of U. reversalis show a consistent geographic pat- 
tern in the USA. Lupinus spp. are the most widespread native hosts; 
Baptisia spp. are hosts in the east, central and south, and Sophora spp. 
are hosts along parts of the south, especially desert areas such as Texas 
and Arizona. In the west, Lupinus is the only reported native host genus 
with the earliest record dating from 1930 in Riverside, California. Intro- 
duced plants in the genera Genista, Spartium, Cytisus, Laburnum, Pip- 
tanthus and Templetonia are reported as hosts throughout the USA. 
These introduced plants are particularly abundant along the western re- 
gions and thus are more frequently reported as hosts of U. reversalis 
than are the relatively less abundant, perennial species of Lupinus. 

The other two families of legumes, Mimosaceae and Caesalpinaceae, 
are reportedly used by one or more species of Uresiphita (Table 1). Ure- 
siphita reversalis was collected from Cassia spp. in the Caesalpinaceae 
(Cassieae) in both California and Florida. Collections of U. reversalis 
from the Mimosaceae are in three tribes: the Ingeae, the Adenthereae, 
and the Acacieae. Species of Acacia (Acacieae) are reported as hosts of 
U. reversalis, U. polygonalis and U. ornithopteralis. Collections of U. re- 
versalis are from Acacia in both California and Texas. Uresiphita polyg- 
onalis were collected from Acacia koa A. Gray in Hawaii, and U. or- 
nithopteralis were collected from an Acacia sp. in Australia. 

Other records include nonleguminous families (Table 2). Uresiphita 
polygonalis was reported from Putterlickia in the South African family 
Celastraceae. Uresiphita polygonalis maorialis was collected from Dis- 
caria (Rhamnaceae) and Chrysanthemum (Asteraceae) in New Zealand. 
And U. ornithopteralis caused heavy damage to willows (Salix) in Aus- 
tralia. Collections and publications of U. reversalis were from 10 to 11 
nonleguminous families, including the Boraginaceae, Buddleiaceae, 
Caprifoliaceae, Geraniaceae, Liliaceae, Myrtaceae, Oleaceae, Rosaceae, 
Rutaceae, Ulmaceae and either the Lauraceae, Myricaceae, or Ericaceae. 
Two or three species are from plants in each of the Caprifoliaceae, 
Oleaceae and Rosaceae. All other families were reported on only one oc- 
casion. Powell (1992) reported two additional families (Taxaceae and Ru- 
biaceae) as possible hosts that I have not included in my collation for the 
following reasons. Both records are from the California Department of 
Food and Agriculture collections. Only pupae were collected from 
Taxus (Taxaceae) and the record or specimen of the collection from 
Gardenia (Rubiaceae) could not be located. Data from Powell’s paper 
were not tabulated since they duplicate information presented here and 
include some questionable data from Bernays and Montllor (1989). 
Host specificity tests on U. reversalis are presented in Leen (1997) and 
clarify this matter. 


146 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Uresiphita reversalis was collected in California on three separate occa- 
sions from barbecue covers originating in Connecticut, Vermont and 
Massachusetts (California Department of Food and Agriculture Records). 
The three collections from barbecue covers exceed the number of times 
larvae were collected from most nonleguminous plants and, obviously, 
barbecue covers are a ‘host’ upon which development is not completed. 


DISCUSSION 


In general, the larval host plants of the genus Uresiphita are confined 
to the quinolizidine-bearing tribes of the Fabaceae. This suggests quino- 
lizidine alkaloids are important to the determination of the host range of 
Uresiphita. The sequestering of quinolizidine alkaloids from G. monspes- 
sulana by U. reversalis was confirmed by Bernays and Montllor (1989) 
and Montllor et al. (1990). Other Uresiphita species also may be found 
to sequester quinolizidine alkaloids since the aposematic coloration, gre- 
garious habits and host plant range are similar among Uresiphita larvae 
(Leen 1992, 1995). One genus, Cytisus, bears quinolizidine alkaloids but 
is not suitable for development of both U. reversalis and U. polygonalis 
(Leen 1992, 1997). Confusion in nomenclature has surely led to erro- 
neous reports on Cytisus and thus all reports remain to be substantiated 
(Leen 1992, 1997). Collections from other genera in tribes of the 
Fabaceae and from the Caesalpiniaceae are questionable because mem- 
bers of these tribes were rejected by Uresiphita and collections are rare. 
However, the collections of three species of Uresiphita from Acacia spp. 
in different localities suggest this may be an accurate report. This is an 
anomalous host plant since Acacia is not known to bear quinolizidine al- 
kaloids although Acacia has been reported to contain other types of alka- 
loids (White 1954, 1957). 

Although some nonleguminous plant families are known to contain 
genera that bear quinolizidine alkaloids (Schwarting 1973, Wink 1992), 
none of the tested genera in these particular families and others were 
acceptable (Leen 1997). Just as the collections from barbecue covers 
are not indicative of host use, most of these collection records are prob- 
ably not indicative of species used by Uresiphita. A few other important 
facts help to discredit these collections as true hosts. Mulvay (1978) 
noted the collection of U. p. maorialis from Chrysanthemum occurred 
because larvae had migrated from their original host, Lupinus. Lonicera 
sempervirens, honeysuckle, is frequently cited as a host plant of U. re- 
versalis. Hedysarum coronarium L. is known as French honeysuckle. 
French honeysuckle may have been a collection host, and the common 
names may have led to confusion. However, both species were rejected 
in the lab and are probably not acceptable hosts under field conditions. 

Species in the Genisteae, Sophoreae, Thermopsidae and Bossiaceeae 


VOLUME 51, NUMBER 2 A, 


are undoubtedly hosts of Uresiphita spp. Further research in regard to 
genera such as Acacia may refute the present conclusions. 


ACKNOWLEDGMENTS 


I thank E. Munroe, K. Hagen and M. Dougherty for reviewing the manuscript and J. 
Santiago-Blay, M. Schaffer, J. Dugdale, J. Brown, and T. Eichlin and for providing infor- 
mation on some of the insect species. I thank P. Kleintjes, S. Tait, B. Des Rochers, J. 
Hamai and J. Andrews for taking care of plants and animals during my absence. Voucher 
specimens of U. reversalis and U. polygonalis are deposited at the Bernice Pauahai Bishop 
Museum, Honolulu, Hawaii. 


LITERATURE CITED 


ANON. 1935. Forest parasite biology. Report. New Zealand State Forest Service: 10. In 
Spiller, D. M. & K. A. J. Wise (1982), A catalogue (1860—1960) of New Zealand in- 
sects and their host plants. Dept. Scientific Industrial Research Bull. 231. Wellington, 
New Zealand. 260 pp. 

ARNETT, JR., R. H. 1985. American insects. Van Nostrand Reinhold Company, New York. 
850 pp. 

Pie A. & C. B. MONTLLOR. 1989. Aposematism of Uresiphita reversalis larvae 
(Lepidoptera). J. Lepid. Soc. 43:261—273. 

CLARKE, J. F. G. 1971. The Lepidoptera of Rapa Island. Smithson. Contr. Zool. 56:1—282. 

. 1986. Pyralidae and Microlepidoptera of the Marquesas Archipelago. Smithson. 
Contr. Zool. 416:1—485. 

COMMON, I. F. B. 1990. Moths of Australia. E. J. Brill, New York. 535 pp. 

CROSSWHITE, C. D. 1985. Damage to mescal bean (Sophora secundiflora) by a Pyralid 
moth (Uresiphita reversalis). Desert Plants 7(1):32. 

FENEMORE, P. G. (ed.) 1982. Plant pests and their control. Butterworths, Wellington, New 
Zealand. 271 pp. ° 

FORBES, W. T. M. 1923. Lepidoptera of New York and neighboring states. Cornell Univ. 
Agric. Expt. Sta. Mem. 68. 729 pp. 

FROGGATT, W. W. 1907. Australian insects. William Brooks and Company Ltd. Sydney, 
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GASKIN, D. E. 1966. The butterflies and common moths of New Zealand. Whitcombe and 
Tombs, Limited. Christchurch, New Zealand. 219 pp. 

GipBs, G. W. 1976. The role of insects in natural terrestrial ecosystems. N. Z. Entomol. 
(O12) 0 a PA 

HANNEMAN, H. 1964. Die tierwelt Deutschlands 50. Teil. kleinschmetterlinge oder Mi- 
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gen (Pyraloidea). 410 pp. Jena. 

HuDson, G. V. 1928. The butterflies and moths of New Zealand. Ferguson and Osborn 
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New Zealand insects and their host plants. Dept. Scientific Industrial Research Bull. 
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KHOTKO, E. I. & R. V. MOLCHANOVvA. 1974. On the morphology of the immature stages of 
some species of the subfamily Pyraustinae (Lepidoptera, Pyralidae). Entomol. Rev. 
Doe) tO5— lb. 

KIMBALL, C. P. 1965. Lepidoptera of Florida. An annotated checklist. Division of Plant 
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LEEN, R. 1992. Not so novel interactions of Uresiphita spp, (Crambidae) and their host 
plants. Unpubl. Ph. D. dissertation. Univ. California, Berkeley. 124 pp. 

. 1995. Biology of Uresiphita reversalis (Guenée) and comparison with U. polygo- 

nalis maorialis (Felder) (Crambidae). J. Lepid. Soc. 49:163—-170. 

. 1997. Host specificity of Uresiphita reversalis (Guenée) (Crambidae). J. Lepid. 

Soc. 51:149—155. 


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LEONARD, M. D. 1926. A list of the insects of New York with a list of the spiders and cer- 
tain other allied groups. Cornell Univ. Agric. Expt. Sta. Mem. 101. 1121 pp: 

Mastro, L. 1990. A study on the natural history of Cytisus (Fabaceae) on Santa Catalina 
Island with an emphasis on biological control. Unpubl. M. Sc. Thesis. California State 
Univ., Long Beach. 78 pp. 

MCKENZIE, H. L. 1933. Observations on the Genista caterpillar. Calif. State Dept. Agric. 
Monthly Bull. 22:410—412. 

MEyRICK, E. 1889. Descriptions of New Zealand Micro-Lepidoptera. Trans. N. Z. Inst. 
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New Zealand insects and their host plants. Dept.Scientific Industrial Research Bull. 
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MILLER, D. 1935. Garden pests in New Zealand. A popular manual for practical garden- 
ers, farmers and schools. Cawthron Institute Monographs 1. 84 pp. 

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alkaloids in the relationship between larvae of Uresiphita reversalis (Lepidoptera: 
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Muvay, R. T. 1978. Biology of the Kowhai Moth Uresiphita polygonalis maorialis. Un- 
publ. M. Sc. Thesis. Univ. Auckland, New Zealand. 30 pp. 

MUNROE, E. 1976. In Dominick, R. B., et al., The moths of America north of Mexico, 
Fasc. 13. 2A, Pyraloidea (in part). E. W. Classey Ltd., England. 150 pp. 

PaLM, E. 1986. Nordeuropas Pyralider: med saerligt henblik pa den danske fauna (Lepi- 
doptera: Pyralidae). Kobenhaven: Fauna Boger. 287 pp. 

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rife. Museo Insula de Ciencias Naturales. Num 2. 157 pp. 

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South Africa. 273 pp. 

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Pac. Entomol. 68:105—121. ; 

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. 1957. Alkaloids of the Leguminosae. Part XXVI: examination of further legumes, 
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Received for publication 29 December 1993; revised and accepted 17 April 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 149-155 


HOST SPECIFICITY OF URESIPHITA REVERSALIS 
(GUENEE) (CRAMBIDAE) 


ROSEMARY LEEN 


United States Department of Agriculture, Forest Service, 
Pacific Southwest Research Station, P. O. Box 236, Volcano, Hawaii 96785, USA 


ABSTRACT. Host specificity tests were conducted on Uresiphita reversalis and to a 
lesser degree on U. polygonalis. First instars of U. reversalis were limited to feeding on 
quinolizidine-bearing tribes of fabaceous legumes. However, U. polygonalis from the Ca- 
nary Islands and U. reversalis both failed to complete development on Cytisus scoparius 
(Genisteae) beyond the second instar. Cytisus scoparius and Cytisus striatus were never 
observed as hosts of U. reversalis in California during the years of this study (1984—1989). 
Host range of U. reversalis encompassed six quinolizidine-bearing tribes of the Fabaceae: 
Genisteae, Sophoreae, Thermopsidae, Bossiaeeae, Podalyreae, and Euchresteae, although 
the latter two tribes have not been reported as hosts in the field. Both native and intro- 
duced species in quinolizidine-bearing tribes will undoubtedly be used by U. reversalis 
when the opportunity arises. 


Additional key words: Pyralidae, Pyraustinae, aposematism, host plant range, 
French broom, quinolizidine alkaloids. 


Uresiphita reversalis (Guenée) expanded its host range from native 
legumes to include several introduced ornamental broom species. Feed- 
ing by U. reversalis on Genista monspessulana (L.) L. Johnson (commonly 
known as French broom or Genista) was first reported to the USDA Agri- 
cultural Research Service, Albany, California, in 1983 when larvae caused 
substantial defoliation of some populations in the San Francisco Bay area. 
These studies were undertaken to determine if U. reversalis might be 
used to control the introduced weedy brooms in California (Leen 1992). 
Unfortunately, plants defoliated in the summer or fall were completely 
refoliated the following spring. Early spring growth of the brooms prior 
to the increase of insect populations also indicated U. reversalis was un- 
likely to be a significant control agent. Studies on the potential host 
range of U. reversalis were completed even though the insect was no 
longer considered a potential, augmentative control agent. 


MATERIALS AND METHODS 


Host acceptance tests of first instars of U. reversalis were conducted 
on insects originating from Alameda County, California, USA and U. 
polygonalis (Denis & Schiffermiiller) originating from Masca, Tenerife, 
Canary Islands, Spain. Uresiphita reversalis was collected from G. mon- 
spessulana, and U. polygonalis was collected from Retama monosperma 
(L.) Boiss. First instars were obtained by collecting and rearing larvae to 
adults and later removing newly laid eggs from foliage before hatching. 
Upon hatching, one or two, and occasionally more, larvae were placed 
on each test plant. An equal number of larvae was used as controls and 


150 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Plants in the Fabaceae accepted by first instars of Uresiphita reversalis. P = 
potted plant tested, C = cutting (excised leaf) tested. 


Hostplant No. insects No. plants P/C 
Genisteae 
Cytisus scoparius (L.) Link 20 20 P 
Cytisus scoparius (Dallimore hybrid) (lilac broom) 10 10 P 
Cytisus striatus (Hill) Rothm. 20 20 C 
Genista lydia Boiss. 2 6 P 
Genista linifolia L. 30 30 P 
Genista monspessulana (L.) L. Johnson 30 30 P 
Genista tinctoria L. 24 24 P 
Genista stenopetala Webb & Berth. Ou 32 P 
Laburnum anagyroides Medik. 26 26 P 
Laburnum alpinum (Mill.) Ber. & J.Presl. 30 30 P 
Lupinus albifrons Benth. 30 30 C 
Lupinus arboreus Sims 30 30 P 
Lupinus chamissonis Eschsch. 30 30 P 
Lupinus luteus L. 20 20 P 
Lupinus succulentus Koch 20 20 P 
Lupinus variicolor Steudel 20 10 C 
Spartium junceum L. 25 DNS) P 
Ulex europaeus L. 20 20 P 
Thermopsidae 
Baptisia australis (L.) R.Br. 30 30 P 
Baptisia lactea (Raf.) Thieret. 30 30 P 
Baptisia tinctoria (L.) Vent. 30 30 P 
Thermopsis rhombifolia Nutt. ex Richards. 30 le P 
Thermopsis macrophylla Hook. & Arn. 30 15 C 
Sophoreae 
Sophora davidii (Franch.) Skeels. 6 3 P 
Sophora secundiflora (Ort.) Lag. ex DC 30 30 P 
Podalyreae 
Podalyria sericea (Andrews) R.Br. 8 4 P 
Euchresteae 
Euchresta Benn. 4 D P 
Vicieae 
Vicia sativa L. (flowers only) 16 40 C 


placed on G. monspessulana cuttings. Development was observed until 
the first instar was completed. Later, tests of U. reversalis and U. polyg- 
onalis on Cytisus scoparius (L.) Link were continued beyond the first 
instar to determine if development could be completed on this species. 
All experiments were conducted on naive larvae under a 16L:8D pho- 
toperiod at 20° C. Developmental tests were conducted on C. scoparius 
because U. reversalis was observed under field conditions to oviposit 
and complete development through the fifth instar on almost all other 


VOLUME 51, NUMBER 2 151 


TABLE 2. Leguminous plants rejected by Uresiphita reversalis larvae. P = potted plant 
tested, C = cutting (excised leaf) tested. 


Hostplant No. insects No. plants P/C Instar 
Fabaceae 
Genisteae 
Cytisus scoparius (L.) Link 20 20 C 2) 
Thermopsidae 
Pickeringia montana Nutt. 4] 1a C ] 
Hedysareae 
Hedysarum coronarium L. 30 30 P ] 
Lespedeza bicolor Turcz. 8 4 P il 
Trifolieae 
Ononis L. 30 30 P IL 
Medicago sativa L. 26 26 P 1 
Trifolium L. 26 26 P Il 
Loteae 
Anthyllis vulneraria L. 30 30 P Ik 
Lotus scoparius (Nutt.) Ottley 25 25 P il 
Vicieae 
Lathyrus latifolius L. 24 2 P 1 
Vicia villosa Roth 9 D5 G 1 
Desmodieae 
Indigofera tinctoria L. 16 8 P Il 
Phaseoleae 
Pueraria lobata (Willd.) Ohwi. 6 3 P il 
Crotalarieae 
Crotalaria capensis Jacq. 8 4 P 1 
Caesalpiniaceae 
Cercidae 
Cercis canadensis L. 20 10 P il 
Cercidium floridum A. Gray 16 8 P 1 
Cassieae 
Ceratonia siliqua L. 6 3 P it 
Mimosaceae 
Ingeae 
Albizia julibrissin Durazz. 8 4 IP IL 
Mimoseae 
Mimosa pudica L. 16 8 P 1 
Leucaena leucocephala (Lam.) DeWit 20 10 P ] 
Acacieae 
Acacia Mill. 10 10 P 1 
Acacia longifolia (Andrews) Willd. 2 6 C IL 


reported hosts in the Genisteae except C. scoparius and Cytisus striatus 
(Hill) Rothm. Again, an equal number of larvae were used as controls 
and placed on G. monspessulana. The plant species used in tests of U. 
reversalis are listed in Tables 1 and 2. First instars of U. polygonalis 
were tested on potted plants of Phaseolus vulgaris L., and an equal 
number of larvae were tested on G. monspessulana. 

Fourth instars of U. reversalis from Alameda County, California, were 


152 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


tested on Lonicera sempervirens L., Convolvulus arvensis L., and Euge- 
nia L. sp. Fourth instars of U. reversalis originating from a population 
near Lake Placid, Florida, and collected from Lupinus diffusus Nutt., 
were also tested on cuttings of L. sempervirens. In each test, one larva 
was tested on each plant and an equal number of larvae were tested on 
G. monspessulana. Both populations were fed G. monspessulana prior to 
testing and observed under the same environmental conditions as above. 

Nearly all potted plant specimens were originally collected as seed 
from locations within California or obtained from a variety of commer- 
cial seed sources and botanical gardens within the USA and abroad. The 
Botanical Garden at the University of California, Berkeley, graciously 
provided many of the seeds from sources outside California. Plants 
grown from seed were fertilized biweekly for the first three months on 
Hoagland’s solution (Hoagland & Arnon 1938). Older plants were then 
fertilized every six to nine months with a timed-release, 17-6-10, fertil- 
izer (Osmocote). Attempts were made to infect test plants with Rhizo- 
bia by inoculating soil with roots infected with Rhizobia from closely re- 
lated plants. A few of the potted plants were obtained by purchasing 
mature plants from nurseries. These potted plants were fertilized with 
Osmocote as above. Tests with cuttings were conducted on plant speci- 
mens obtained from localities within California and initiated within 48 
hours from the time of collection. 


RESULTS 


First instars of U. reversalis from California accepted 27 plant species 
from five tribes (Genisteae, Thermopsidae, Sophoreae, Podalyreae, and 
Euchresteae) in the Fabaceae (Table 1). All accepted tribes are well 
represented by species bearing quinolizidine alkaloids (Wink 1992) with 
a few exceptions. Pickeringia montana Nutt., in the Thermopsidae, is 
not known to contain quinolizidine alkaloids and was rejected by U. re- 
versalis (Table 2). Flowers, but not leaves, of Vicia sativa L. in the Vi- 
cieae were accepted by U. reversalis. Neither this species nor the tribe 
are reported to contain quinolizidine alkaloids. The foliage of V. sativa 
and the foliage and flowers of Vicia villosa were both unacceptable to U. 
reversalis (Table 2). 

Fourteen species from eight tribes (Thermopsidae, Hedysareae, Tri- 
folieae, Loteae, Vicieae, Desmodieae, Phaseoleae and Crotalarieae) in 
the Fabaceae were rejected by first instars of U. reversalis (Table 2). 
Eight species from five tribes of nonfabaceous legumes were also re- 
jected by first instars (Table 3). Thirty two species in 12 nonleguminous 
families were rejected by first instars, and three species in three families 
were rejected by fourth instars (Table 2.) Some of these rejected fami- 
lies (e.g., Ranunculaceae, Scrophulariaceae) were chosen for testing be- 


VOLUME 51, NUMBER 2 153 


TABLE 3. Non-leguminous plants rejected by Uresiphita reversalis larvae. P = potted 
plant tested, C = cutting (excised leaf) tested. 


Hostplant No. insects No. plants P/C Instar 

Caprifoliaceae 

Lonicera japonica Thumb. 19 4 P 1 

Lonicera hispidula Doug. 45 5 P 1 

Lonicera sempervirens L. 40 20 P 1 

Lonicera sempervirens L. 15 15 P 4 

Sambucus mexicana C. Presl. 8 4 P 1 

Symphoricarpus albus (L.) S.F.Blake 5 5 P i 
Asteraceae 

Arctium minus (Hill) Bernh. 24 24 P i 

Calendula officinalis L. 20 20 P 1 

Centaurea cyanus L. 24 24 P 1 

Centaurea diffusa Lam. 48 48 P 1 

Centaurea maculosa Lam. 24 24 P Il 

Chrysanthemum leucanthemum L. 30 30 P 1 

Chrysanthemum parthenium (L.) Bernh. 30 30 P iL 

Helianthus tuberosus L. 40 20 P 1 

[satis tinctorius L. oo, 16 I? 1 

Santolina chamaecyparissus L. 19 3) P 1 

Serratula radiata (Waldst. & Kit.) Bieb. 24 24 P IL 

Silene italica (L.) Pers. 20 20 P il 

Tagetes erecta L. 8 4 P 1 
Euphorbiaceae 

Euphorbia esula L. 20 20 P 1 
Convolvulaceae 

Convolvulus arvensis L. I5 25 P 1 

Convolvulus arvensis L. 20 20 P 4 
Papaveraceae 

Eschscholzia californica Cham. 30 30 P Th 

Papaver orientale L. 30 30 P 1 

Papaver somniferum L. 46 46 P 1 
Ranunculaceae 

Cimicifuga racemosa (L.) Nutt. 20 Ht P 1 

Aconitum napellus L. 20 1 P 1 
Malvaceae 

Malva alcea L. 24 24 P 1 
Scrophulariaceae 

Antirrhinum majus L. 20 20 1) 1 
Plantaginaceae 

Plantago lanceolata L. 24 24 12 1 
Brassicaceae 

Brassica oleracea L. 20 20 P iL 
Lamiaceae 

Mentha aquatica L. 24 24 P 1 
Myrtaceae 

Eugenia L. 105) it C 4 
Boraginaceae 


Ehretia anacua (Teran & Berl.) I.M. Johnson 45 30 P i 


154 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


cause they are reported to contain species bearing quinolizidine alka- 
loids. Several of the rejected plant species (L. sempervirens, Ehretia 
anacua (Teran & Berl.) I. M. Johnston and Eugenia) were reported as 
hosts of U. reversalis. 

Although U. reversalis completed development on C. scoparius and 
C. striatus through the first instar (Table 1), larvae did not complete de- 
velopment beyond the second instar on C. scoparius (Table 2). Ure- 
siphita polygonalis did not complete development beyond the second 
instar on C. scoparius (n = 20 potted plants tested) or beyond the first 
instar on P. vulgaris (n = 22 potted plants tested). Fourth instars of U. 
reversalis from California did not feed upon nonleguminous plants 
(Table 3). All larvae died before molting or pupating. The Floridean 
population of U. reversalis also refused to accept L. sempervirens (n = 
15 cuttings tested). Most of the rejected plants are not known to bear 
quinolizidine alkaloids. Control larvae rarely died or failed to complete 
development on G. monspessulana. Observed deaths were attributed to 
handling problems rather than to the control plants and are therefore 
not tabulated. 


DISCUSSION 


There are inconsistencies among reported hosts and host acceptance 
tests of Uresiphita. Although C. scoparius is a reported host for several 
species of Uresiphita, the accuracy of such reports is questionable for 
several reasons. First, rejection of C. scoparius by both U. reversalis and 
U. polygonalis indicates this species could not support these larvae 
through complete development. Second, C. scoparius is frequently con- 
fused with G. monspessulana by collectors in California. Insect speci- 
mens are thus labelled incorrectly with records of Scotch broom, 
Cytisus or C. scoparius, as the host plant. Third, G. monspessulana was 
classified as Cytisus monspessulanus L. in several floras. Inaccurate 
records for other species of Uresiphita in regard to Cytisus may also exist. 
The rejection of C. scoparius by U. reversalis and U. polygonalis does not 
exclude the possibility that other species of Uresiphita use Cytisus and are 
able to complete development. An explanation as to why C. scoparius is 
apparently the only rejected species in the tribe Genisteae cannot 
presently be offered. Tests on C. striatus were not conducted beyond 
the first instar for U. reversalis (Table 4). Larvae may be unable to com- 
plete development beyond the second instar on other species of Cytisus. 

Bernays and Montllor (1989), citing my preliminary host plant data 
for first instars, reported that feeding does not occur upon Pickeringia, 
Trifolium, Vicia, and Medicago and that extensive feeding occurs on C. 
scoparius, C. striatus, L. arboreus, and G. monspessulana. They also 
stated that development cannot be completed upon Laburnum or Ulex. 


VOLUME 51, NUMBER 2 WEE 


Only the information on L. arboreus, G. monspessulana, Pickeringia 
montana (a monotypic genus), Trifolium, and Medicago is accurate. 

Although some nonleguminous plant families are known to contain 
genera that bear quinolizidine alkaloids (Schwarting 1973, Wink 1992), 
none of the tested genera in these particular families and others were 
acceptable. Most of these collection records are probably not indicative 
of species used by Uresiphita. 

Two genera (Adenostoma, Rosa) in the Rosaceae have been reported 
as hosts of U. reversalis. The collection and rearing of larvae from 
Adenostoma fasciculatum Hook. & Arn. was from a location where other 
probable hosts are not present (the old lighthouse at Point Loma, Cali- 
fornia) and thus is assumed accurate. First instars of U. reversalis did 
not complete development on A. fasciculatum in the lab. Two explana- 
tions are offered for the conflicting collection record and laboratory re- 
sults. One, A. fasciculatum may be an acceptable host for later instars if 
U. reversalis was transferred (e.g., by humans) onto Adenostoma. Two, 
the source of tests plants of A. fasciculatum was central California rather 
than southern California where the insect was collected. Host plant vari- 
ation may explain the laboratory rejection of A. fasciculatum. 

Larval hosts of Uresiphita spp. are primarily limited to quinolizidine- 
bearing tribes of the Fabaceae (Leen 1992 1997) and larval hosts of U. 
reversalis are similarly limited in range. Native hosts come from three 
tribes: Genisteae, Sophoreae, and Thermopsidae. However, host speci- 
ficity tests, collections, and publications indicate additional species bear- 
ing these alkaloids will be utilized when the opportunity arises. 


ACKNOWLEDGMENTS 


I thank E. Munroe, K. Hagen, and M. Dougherty for reviewing the manuscript and P. 
Kleintjes, S. Tait, B. Des Rochers, J. Hamai, and J. Andrews for taking care of plants and 
animals during my absence. Voucher specimens of U. reversalis and U. polygonalis are de- 
posited at the Bernice Pauahai Bishop Museum, Honolulu, Hawaii. 


LITERATURE CITED 


BERNAYS, E. A. & C. B. MONTLLOR. 1989. Aposematism of Uresiphita reversalis larvae 
(Lepidoptera). J. Lepid. Soc. 43:261—273. 

HOAGLAND, D. R. & D. I. ARNON. 1938. The water-culture method for growing plants 
without soil. Univ. Calif. Circular 347. 39 pp. 

LEEN, R. 1992. Not so novel interactions of Uresiphita spp, (Crambidae) and their host 
plants. Unpubl. Ph.D. dissertation, Univ. California, Berkeley. 124 pp. 

—. 1997. Larval hosts of Uresiphita Hiibner (Crambidae). J. Lepid. Soc. 51:139-148. 

SCHWARTING, A. E. 1973. The quinolizidine alkaloids. Nobel 25 chemistry in botanical 
classification, pp. 205—210. 

WINK, M. 1992. The role of quinolizidine alkaloids in plant-insect interactions, pp. 
131-166. In Bernays, E. A. (ed.), Insect-plant interactions. Volume IV. CRC Press, 
Boca Raton, Florida. 


Received for publication 29 December 1993; revised and accepted 17 April 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 156-175 


DISTRIBUTION AND PHENOLOGIES OF 
LOUISIANA SPHINGIDAE 


VERNON ANTOINE BROU, JR. 
AND 


CHARLOTTE DOZAR BROU 
74320 Jack Loyd Road, Abita Springs, Louisiana 70420, USA 


ABSTRACT. The abundance, distribution, and flight periods for 55 species of 
Louisiana Sphingidae are presented, including prior literature records and new collecting 
data for 44 species taken over a 26-year period (1970 through 1995). Information is pro- 
vided on the number of annual broods for 36 species, and dates of capture are plotted as 
one-year and composite graphs for 30 species. 


Additional key words: _ bait traps, hawkmoths, light traps, sphinx moths, voltinism. 


The first treatment of the family Sphingidae in Louisiana was pre- 
sented by von Reizenstein (1863), who reported 33 species from the 
vicinity of New Orleans. Later, von Reizenstein (1881) and Ottolengui 
(1894) each tallied one additional species for the state, and subsequent 
published works during this century have gradually expanded the total 
by another dozen species (see Rothschild & Jordan 1903, Hine 1906, 
Clark 1917, Draudt 1931, Jung 1950, Merkl & Pfrimmer 1955, Pfrim- 
mer 1957, Brou 1980, Covell 1984, Brou 1994). For some time, we have 
been monitoring and collecting adult Louisiana Sphingidae in order to 
produce a comprehensive state list and examine voltinism and variation 
in the abundance of adults from brood to brood. In the present paper, 
we discuss the results from 26 years of sampling, 1970 through 1995. 


MATERIALS AND METHODS 


We used ultraviolet light traps and fermenting bait traps to attract 
sphingids, logging approximately 416,000 light trap hours and 633,000 
bait trap hours from 1970 through 1995. Occasional sampling was done 
using hand nets, flight traps, and pitfall traps. Many different light trap 
designs were used, but generally the traps employed lamps with adja- 
cent baffles mounted over a funnel (see Brou 1992a, 1992b for details). 
Most of the lamps were black lights, ranging from 15 to 1000 watts, used 
singly, or in various combinations. Light traps varied from 60 to 3500 
watts each, though most were in the 250 to 600 watt range. As many as 
six light traps were operated dusk to dawn, irrespective of climatic con- 
ditions, using photoelectric controls. Up to eight bait traps were oper- 
ated year-round during 1984—1995. 

Brood numbers were estimated by examining yearly graphs of capture 
totals plotted against sampling date for individual sphingid species. Ap- 


VOLUME 51, NUMBER 2 WSR. 


proximately 2000 such graphs were prepared and studied, yielding data 
sufficient to estimate the number of annual broods for 36 species. Rep- 
resentative single-year graphs and composite-year graphs are presented 
in Figs. 3-52 (see Results and Species Accounts for discussion). 

Specimens retained during this study are deposited at several institu- 
tional and private collections, the largest numbers of specimens being in 
the Florida State Collection of Arthropods (Gainesville), Louisiana State 
University (Baton Rouge) and in the collection of the senior author. 
Most specimens were from nine locations that were monitored on a fre- 
quent or continuous basis. These were, in decreasing order of sampling 
intensity: St. Tammany Parish, Sec. 24, T6, SR12E, 6.8 km NE Abita 
Springs; St. John the Baptist Parish, Edgard; Iberville Parish, Sunshine; 
Lafourche Parish, Cut Off; West Feliciana Parish, Sec. 63 and Sec. 76, 
T1S, R3W, 3.2 km NE Turnbull/Weyanoke; Ascension Parish, Prairie- 
ville; Tangipahoa Parish, Fluker; Natchitoches Parish, Kisatchie National 
Forest; Orleans Parish, New Orleans. Nomenclature follows Hodges 
(1983) with minor modifications. 


RESULTS 


General Trends. A total of 71,836 specimens of 55 species of Sphin- 
gidae was sampled from 43 of 64 Louisiana parishes (Fig. 1). Of the re- 
maining 21 parishes, some were not visited, and a few yielded no Sphin- 
gidae. The greatest number of species (40) was recorded from St. 
Tammany Parish. Distribution maps for each of the 55 species are pre- 
sented in Figs. 2.1—-2.55, using data from our study supplemented by 
those few prior literature records for which accurate locality information 
could be determined. 

Table 1 lists the monthly sampling totals from our study for each of 
the 55 species. Over 96 percent of the specimens came to light or bait 
during the period March to September. All 55 of the sphingid species 
that we sampled came to ultraviolet light traps (including Hemaris 
thysbe (F.), H. diffinis (Bdv.), and Amphion floridensis B. P. Clark). Da- 
rapsa myron (Cram.) was the most common species, accounting for over 
26 percent of the total. Species more often taken in fermenting bait 
traps included Sphinx kalmiae Neum., Enyo lugubris (L.), Sphecodina ab- 
bottii (Swainson), A. floridensis, D. myron, and Darapsa pholus (Cram.) 
(Platt (1969) reported collecting some of these same species at fruit 
bait). Several specimens each of species not generally known to be at- 
tracted to fermenting bait were taken by this method, including Laothoe 
juglandis (J. E. Smith), H. diffinis, Darapsa versicolor (Harr.), and Xylo- 
phones tersa (L.). Several Agrius cingulata (F.) and Amphion floridensis 
were captured in pitfall traps baited with a mixture of feces, water, and 


ethylene glycol. 


158 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Number of Sphingidae species recorded for each Louisiana parish, from 
sampling during 1970-1995. 


A total of 47 of the 55 species had been recorded previously from 
Louisiana, and 8 are reported here as new state records (B in Table 1). 
We failed to locate six species recorded by von Reizenstein (1863, 1881) 
(V in Table 1), two recorded by Jung (1950) (J in Table 1), and one 
recorded by Ottolengui (1894) (O in Table 1). We have been unable to 
locate specimens from the literature reports tabulated earlier, and Jung 
(pers. comm.) indicated that specimens taken during his investigation 
no longer exist. Two species, Sphinx leucophaeta Clem. and Sphinx 
chersis (Hbn.), reported by von Reizenstein (1863) seem questionable, 
although these species are known from one or more adjoining states. 
These records may actually refer to Sphinx franckii Neum., which was 
not recognized and named until 30 years after von Reizenstein’s publi- 
cation. Ottolengui’s (1894) report of Eumorpha licaon (Cram.) likely was 


159 


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VOLUME 51, NUMBER 2 163 


Eumorpha intermedia (B. P. Clark), a similar, smaller species, not de- 
scribed until 23 years later. 

Among sphingid surveys from adjoining states, Freeman (1938) 
recorded 32 species from Arkansas, and only one of his species, Sphinx 
gordius Cramer, remains unrecorded from Louisiana (Riotte (1980) sug- 
gested that this record was probably Sphinx poecila Stephens). All the 
sphingids in the following three surveys are known from Louisiana: the 
28 species listed by Selman and Barton (1971) from northeastern 
Arkansas; the 26 species listed by Neck (1991) from Walker County, 
Texas; and the 24 species listed by Taylor and Taylor (1965) from the 
Gulf Coast of Mississippi. 

Annual Brood Patterns. We were able to estimate the number of 
annual broods in Louisiana for 36 of 55 species of Sphingidae (see Spe- 
cies Summaries). Nearly all of our findings differ from previously pub- 
lished sphingid voltinism in other states (e.g., Beutenmuller 1895, 
Hodges 1971, Covell 1984, Heitzman 1987), with Louisiana’s southerly 
location generally promoting additional broods. For many of the multi- 
brooded species, the interval between the first brood peak and the sec- 
ond brood peak in any given year proved to be sometimes two times 
greater than the intervals between the remaining brood peaks; these 
subsequent intervals were usually consistent, or nearly so, throughout 
the remainder of the year. This initial nonconforming brood interval is 
of different duration depending on the species. The initial spring broods 
of some species can also be quite protracted, likely influenced at least in 
part by unpredictable spring climatic influences upon both the moths 
and their foodplants. The initial brood peaks varied by a month or more 
from one year to another. Variability in initial brood emergence appears 
to affect the timing of subsequent broods, but the magnitude of the ef- 
fect differs in any given year. 

Certain annual broods in some species also tend to be consistently 
small or large, and these relative brood sizes tend to repeat from year to 
year. For example, the fourth annual brood of Darapsa myron is typi- 
cally the smallest, and is bordered by the two largest broods (three and 
five) (Figs. 29, 33). For Sphecodina abbottii, the second and especially 
the fourth broods are reduced (the fourth may in fact represent a fre- 
quently observed partial brood). The composite-year graph for S. abbot- 
tii (Fig. 25) masks these small broods, but the trends are more apparent 
when individual years are examined (cf. Fig. 34). A similar pattern can 
be seen in D. pholus, in which broods occur at approximately monthly 
intervals. Broods one, three, five, and six are usually more populous than 
broods two, four, and seven. On the composite-year graph (Fig. 30), 
broods five through seven merge together and give the impression of a 
single, final brood (cf. Fig. 35). Why some multibrooded species have 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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166 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


. A. ciggulata 


4.4 sexta 


_ M. yesminearum 
, [2 hyloeus 


. C amynator 


. C undulosa 


10. Z cupressf 
1l. & plebeys 


12. S. ta/mise 


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 


Fics. 3-12. Phenologies of Louisana Sphingidae: composite-year graphs for individual 
species, data summed from 1970-1995. Specimens sampled (n) and number of specimens 
represented by entire vertical axis (y) given at right. 3, Agrius cingulata; 4, Manduca sexta; 
5, M. rustica; 6, M. jasminearum, 7, Dolba hyloeus; 8, Ceratomia amyntor; 9, C. undu- 
losa; 10, Isoparce cupressi; 11, Paratrea plebeja; 12, Sphinx kalmiae. 


VOLUME 51, NUMBER 2 167 


13. L. coaierarugt 


14. Z. phacobrachycerous ; 
15. S semarcensis 

16. F exeaocatus’ 

18. 2 asty/us 

19. L. juglandis | 


20. E lugubris 


21. H. thysbe , ‘n=236 
. , y=50 


22. & paadorus ' 


Jan Feb Mar Apr May Jun Jul Auq Sep Oct Nov Dec 


Fics. 13-22. Phenologies of Louisana Sphingidae: composite-year graphs for individ- 
ual species, data summed from 1970-1995. Specimens sampled (n) and number of speci- 
mens represented by entire vertical axis (y) given at right. 13, Lapara coniferarum; 14, L. 
phaeobrachycerous; 15, Smerinthus jamaicensis; 16, Paonias excaecatus; 17, P. myops; 
18, P. astylus; 19, Laothoe juglandis; 20, Enyo lugubris; 21, Hemaris thysbe; 22, Eumor- 
pha pandorus. 


168 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


. & intermedia 


_ & fascista 
. abbott! 
_B inscriote : 
. A. Hloritonsks. 


. D. versicolor ' 


. D pholus 


. . Macata 


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 


Fics. 23-32. Phenologies of Louisana Sphingidae: composite-year graphs for individ- 
ual species, data summed from 1970-1995. Specimens sampled (n) and number of speci- 
mens represented by entire vertical axis (y) given at right. 23, Eumorpha intermedia; 24, 
E. fasciata; 25, Sphecodina abbottii; 26, Deidamia inscripta; 27, Amphion floridensis; 28, 
Darapsa versicolor; 29, D. myron; 30, D. pholus; 31, Xylophanes tersa; 32, Hyles lineata. 


VOLUME 51, NUMBER 2 169 


reduced population sizes for certain broods remains unclear. No doubt 
climatic extremes (e.g., rainfall, drought) and biological influences (e.g., 
predators, parasites) play a role, but we neither systematically studied 
nor found obvious correlations between these factors and observed 
brood sizes and timings. 

The approximate 30-day brood cycles that were exhibited by many 
species are not sampling artifacts related to the lunar cycle, as species 
attracted to fermenting bait showed the same cyclical patterns as those 
attracted to light. A good example is E. lugubris in 1991 (Fig. 36). This 
species is attracted to both light and bait. The initial 1991 brood oc- 
curred in early to mid July, roughly coinciding with a new moon. Broods 
two through five peaked at about 28-day intervals beginning in early 
September, and these subsequent brood peaks did not coincide with ei- 
ther new or full moons (persistent cold weather during December 1991 
prevented collection of sixth brood specimens). Similarly, L. phaeo- 
brachycerous Brou in 1991 (Fig. 37) showed brood peaks not correlated 
with lunar phase. Few specimens of the initial brood were collected in 
early May, as is normally true for this species, and the remaining four 
brood peaks occured at about 30-day intervals between new and full 
moons, beginning in early June. 

For species often seen only at low numbers, representative specimens 
for each brood were not observed in some years. An example is Isoparce 
cupressi. During 1991, only 14 specimens of the initial brood were taken 
(Fig. 40); a single specimen for the second brood; two specimens for the 
third brood; and 16 specimens for the fourth brood. In 1980 (Fig. 41), 
no specimens were collected at the usual emergence time of the second 
brood, though there were specimens representing broods three, four, 
and five. In 1978 (Fig. 42), the first and fifth broods were each repre- 
sented by single specimens, whereas broods two, three, and four were 
represented by multiple specimens. In 1973 (Fig. 43), only broods two, 
three, and four were represented. | 


SPECIES SUMMARIES 


Agrius cingulata (F.) (Fig. 3): seven broods, first peaking late April to early May; peaks 
two through seven occur at approximately 30-day intervals, beginning early to mid-June; 
occasional December specimens may indicate partial emergence of an eighth brood. 

Manduca sexta (L.) (Figs. 4, 44): five broods peaking at approximately 30-day intervals, 
beginning at the end of April; occasional October specimens may indicate partial emer- 
gence of a sixth brood; previously reported by Beutenmuller (1895) as double-brooded 
near New York City, and by Heitzman (1987) as having two or more broods in Missouri. 

Manduca quinquemaculata (Haw.): five broods, first peaking approximately mid April; 
peaks two through five occur at approximately 30-day intervals, beginning early June; oc- 
casional October specimens may indicate partial emergence of a sixth brood; previously 
reported by Beutenmuller (1895) as double-brooded near New York City, and by Heitz- 
man (1987) as having two or more broods in Missouri. 


170 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


33. D pes 1985 : : : : : : : : : n=1772 
, 5 ' ' ' ' a wens 

is abbott! 1987 

ny 2) pholus 1989 

5 (4, ligubri io 

5, L phacobrachycerous 1991 

» Lb phscobrackroerous 1990 

Lh phscobrachycerous 1989 


LL cupressi 1991 


. £ cupresst 1980 


. L cupressi 1978 


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 


Fics. 33-42. Phenologies of Louisana Sphingidae: single-year graphs for individual 
species. Specimens sampled (n) and number of specimens represented by entire vertical 
axis (y) given at right. Full moons on Figs 36, 37 indicated by open circles (see text for 
elaboration). 33, Darapsa myron, 1985; 34, Sphecodina abbottii, 1987; 35, Darapsa pho- 
lus, 1989; 36, Enyo lugubris, 1991; 37, Lapara phaeobrachycerous, 1991; 38, L. phaeo- 
brachycerous, 1990; 39, L. phaeobrachycerous, 1989; 40, Isoparce cupressi, 1991; 41, L 
cupressi, 1980; 42, I. cupressi, 1978. 


VOLUME 51, NUMBER 2 eal 


By 4 cipro 1973 

. Wt. sexte 1984 

ou 4 unduloss 1 991 

as tindulose 1983 

2. contforarum 1989 : 
Bey & coniferarum 1990 : 
. A. Horidensie 1984 


. A. floridensis. 1985 


_ A, forsdeasis' 1990 


. H. Maeata 1991 


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 


Fics. 43-52. Phenologies of Louisana Sphingidae: single-year graphs for individual 
species. Specimens sampled (n) and number of specimens represented by entire vertical 
axis (y) given at right. 43, Isoparce cupressi, 1973; 44, Manduca sexta, 1981; 45, Cerato- 
mia undulosa, 1991; 46, C. undulosa, 1983; 47, Lapara coniferarum, 1989; 48, L. conifer- 
arum, 1990; 49, Amphion floridensis, 1984; 50, A. floridensis, 1985; 51, A. floridensis, 
1990; 52, Hyles lineata, 1991. 


172 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Manduca rustica (F.) (Fig. 5): four broods, first peaking at the end of May; peaks two 
through four occur at approximately 30-day intervals, beginning early July. 

Manduca jasminearum (Guer.) (Fig. 6): two broods, peaking early June and mid Au- 
gust; similar brood times occur for S. franckii; previously reported by Beutenmuller (1895) 
as probably double-brooded near New York City. 

Dolba hyloeus (Drury) (Fig. 7): six broods, first peaking in early April; peaks two 
through six occur at approximately 30-day intervals, beginning mid-May; occasional Octo- 
ber specimens may indicate partial emergence of a seventh brood; previously reported by 
Rowley (1899) as probably double-brooded in Missouri. 

Ceratomia amyntor (Geyer) (Fig. 8): five broods, first peaking in early April; peaks two 
through five occur at approximately 30-day intervals, beginning mid-May; previously re- 
ported by Hodges (1971) as having two broods in the south, by Covell (1984) as having 
two broods, and by Heitzman (1987) as having two broods in Missouri. 

Ceratomia undulosa (Wlk.) (Figs. 9, 45, 46): six broods, first peaking in early April; 
peaks two through six occur at approximately 30-day intervals; initial brood emergence 
varying by two weeks from year to year; previously reported by Beutenmuller (1895) as 
double-brooded near New York City, and by Hodges (1971) and Covell (1984) as having 
two broods. 

Ceratomia catalpae (Bdv.): five broods, first peaking in mid April; peaks two through 
five occur at approximately 35-day intervals, beginning late May; previously reported by 
Hodges (1971) and Covell (1984) as having two broods. 

Ceratomia hageni Grt.: four or more broods; additional records are needed; previously 
reported by Hodges (1971) and Covell (1984) as having two broods, and by Heitzman 
(1987) as having three broods in Missouri. 

Isoparce cupressi (Bdv.) (Figs. 10, 40—43): usually four broods, protracted initial brood 
usually peaking third week of March; peaks two through four occur at approximately 50- 
day intervals, beginning late May; initial emergence peak varying by three weeks from year 
to year; affecting emergence time of subsequent broods (and probably why in some years 
there are specimens representing five broods); previously reported by Covell (1984) as 
having two broods. 

Paratrea plebeja (F.) (Fig. 11): six broods, first peaking early to mid April; peaks two 
through six occur at approximately 30-day intervals, beginning five weeks later; previously 
reported by Beutenmuller (1895) as double-brooded near New York City, by Hodges 
(1971) as having two broods in the south, and by Holland (1903), Rothschild & Jordan 
(1903), and Covell (1984) as having two broods. 

Sphinx franckii Neum.: two broods, peaking approximately mid June and mid August; 
previously reported by Hodges (1971) and Covell (1984) as having one brood and a partial 
second. 

Sphinx kalmiae J. E. Smith (Fig. 12): six broods, first peaking early to mid April; re- 
maining peaks at approximately 30-day intervals; previously reported by Beutenmuller 
(1895) as double-brooded near New York City, and by Hodges (1971) as probably having 
two broods. 

Lapara coniferarum (J. E. Smith) (Figs. 13, 47, 48): five broods, first peaking early to 
mid April (see Brou 1994); peaks two through five occur at approximately 30-day intervals, 
beginning mid-June; previously reported by Koebele (1881) as having at least two broods 
in the southern United States, and by Riotte (1972) as having two distinct flight periods in 
the south coastal states. 

Lapara phaeobrachycerous Brou (Figs. 14, 37—39): five broods, first peaking about mid 
May (see Brou 1994); remaining peaks occur at approximately 30-day intervals. 

Smerinthus jamaicensis (Drury) (Fig. 15): five or more broods, protracted initial brood 
peak approximately mid March; peaks two through four occur at approximately 45-day in- 
tervals, beginning early-June; November specimens may indicate a partial sixth brood; 
previously reported by Beutenmuller (1895) as being double-brooded near New York City, 
and by Heitzman (1987) as being multibrooded in Missouri. 

Paonias excaecatus (J. E. Smith) (Fig. 16): four broods, first peaking late March; peaks 
two through four occur at approximately 45-day intervals, beginning early June; previously 
reported by Beutenmuller (1895) as double-brooded near New York City, by Rowley (1898) 


VOLUME 51, NUMBER 2 7S 


as double-brooded in Missouri, by Hodges (1971) as having two broods in Florida, by Covell 
(1984) as having three broods, and by Heitzman (1987) as having several broods in Missouri. 

Paonias myops (J. E. Smith) (Fig. 17): four broods, first peaking late March; peaks two 
through four occur at approximately 50-day intervals, beginning early June; previously re- 
ported by Beutenmuller (1895) as probably double-brooded near New York City, by 
Hodges (1971) as seemingly single-brooded, and by Heitzman (1987) as having multiple 
broods in Missouri. 

Paonias astylus (Drury) (Fig. 18): four broods; limited data indicate it may have broods 
similar to other members of the genus; previously reported by Hodges (1971) as having 
two broods in Florida, and by Covell (1984) as having two broods. 

Laothoe juglandis (J. E. Smith) (Fig. 19): four broods, peaking at approximately 45-day in- 
tervals, beginning late April; previously reported by Beutenmuller (1895) as double-brooded 
near New York City, by Hodges (1971) as having two broods in the south, by Covell (1984) 
as having three broods, and by Heitzman (1987) as having several broods in Missouri. 

Pachysphinx modesta (Harr.): five broods, first peaking late March; peaks two through 
five occur at approximately 30-day intervals, beginning mid-May; previously reported by 
Beutenmuller (1895) as probably double-brooded near New York City, by Hodges (1971) 
as having two broods in Arkansas, Kansas, and perhaps Missouri, by Covell (1984) as hav- 
ing three broods, and by Heitzman (1987) as being multibrooded in Missouri. 

Enyo lugubris (L.) (Figs. 20, 36): usually six broods, first peaking about mid July; peaks 
two through six occur at approximately 30-day intervals, beginning early September; 
broods five, six, and occasionally seven affected by cold weather during some years; previ- 
ously reported by Holland (1903) as having two broods in Florida. 

Hemaris thysbe (F.) (Fig. 21): six broods, first peaking end of March, and at approxi- 
mately 30-day intervals; previously reported by Beutenmuller (1895) as double-brooded 
near New York City, by Rowley (1899) as double-brooded in Missouri, by Hodges (1971) 
as having two broods in the south, by Covell (1984) as having two broods, and by Heitz- 
man (1987) as having three broods in Missouri. 

Hemaris diffinis (Bdv.): four broods, first peaking mid April, and at approximately 50- 
day intervals; previously reported by Rowley (1899) as double-brooded in Missouri, by 
Hodges (1971) as double-brooded in the northern United States, and by Covell (1984) as 
having two broods. 

Eumorpha pandorus (Hbn.) (Fig. 22): four broods, first peaking about mid May; peaks 
two through four occur at 30-day intervals, beginning early July; previously reported by 
Beutenmuller (1895) as double-brooded near New York City, and by Rowley (1899) as 
double brooded in Missouri. 

Eumorpha intermedia (B. P. Clark) (Fig. 23): four broods, first peaking about mid May; 
peaks two through four occur at approximately 30-day intervals beginning late June. 

Eumorpha fasciatus (Sulz.) (Fig. 24): six or more broods, first peaking in early May, and 
at approximately 30-day intervals; initial brood emergence varying by two weeks in any 
given year; November and December specimens may indicate partial emergence of sev- 
enth and eighth broods; previously reported by Hodges (1971) as having two broods in 
South Carolina, and Covell (1984) as having two broods. 

Sphecodina abbottii (Swainson) (Figs. 25, 34): three or four broods, first peaking end of 
March, and at approximately 45-day intervals; initial brood emergence varying by more 
than two weeks in any given year; broods two and four occur at low numbers; previously 
reported by Heitzman (1987) as having two broods in Missouri. 

Deidamia inscripta (Harr.) (Fig. 26): one brood, peaking at the end of March; previ- 
ously reported by Beutenmuller (1895) as probably double-brooded near New York City, 
and by Hodges (1971) as having one brood. 

Amphion floridensis B. P. Clark (Figs. 27, 49-51): six broods, first peaking end of 
March, and at approximately 25-day intervals; initial brood emergence varying by two 
weeks in any given year; previously reported by Hodges (1971) as multiple-brooded in the 
south, and by Covell (1984) as having two broods. 

Darapsa versicolor (Cram.) (Fig. 28): five or more broods, peaking at approximately 30- 
day intervals; additional records are needed; previously reported by Beutenmuller (1895) 
as double-brooded near New York City, and by Forbes (1948) as having two broods. 


174 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Darapsa myron (Cram.) (Figs. 29, 33): five or more broods, first peaking early to mid 
April, and at approximately 30-day intervals; initial brood emergence varying by more than 
two weeks in any given year; September and October specimens may represent partial 
emergence of sixth and seventh broods; previously reported by Beutenmuller (1895) as 
double-brooded near New York City, by Hodges (1971) as double-brooded in New York 
and South Carolina, and by Covell (1984) as having two broods. 

Darapsa pholus (Cram.) (Figs. 30, 35): seven broods, first peaking end of March, and at 
approximately 30-day intervals; initial brood emergence varying by two weeks in any given 
year; second, fourth, and seventh broods usually at low numbers; November specimens 
may represent partial emergence of an eighth brood; previously reported by Beutenmuller 
(1895) as double-brooded near New York City, by Rowley (1898) as having two broods in 
Missouri, and by Lutz (1948), Hodges (1971), and Covell (1984) as having two broods. 

Xylophones tersa (L.) (Fig. 31): six or more broods, first peak variable, usually at the 
start of May; peaks two through six occur at approximately 30-day intervals, beginning 
mid-June; occasional late year specimens appear, probably representing partially emergent 
subsequent brood(s). 

Hyles lineata (F.) (Figs. 32, 52): eight or nine broods, variable first peak late February 
to early March, and at approximately 30-day intervals; previously reported by Beuten- 
muller (1895) as double-brooded near New York City. 


ACKNOWLEDGMENTS 


We thank the following individuals who supplied specimens or records, or aided in the 
successful completion of this project: Gary Adams, Linda and Phil Auld, Howard D. 
Baggett, April R. Brou, Joan B. Chapin, Kevin J. Cunningham, Douglas C. Ferguson, H. 
Avery Freeman, Lawrence F. Gall, Michael L. Israel, Rodney Jung, Jonathan Kemp, Rick 
Kergosien, Michael T. Lefort, Zack Lemann, Michael Lockwood, Bryant Mather, Eric H. 
Metzler, Eric L. Quinter, Gayle Strickland, Howard V. Weems Jr., and Frances C. Welden. 


LITERATURE CITED 


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miles of New York City. Bull. Am. Mus. Nat. Hist. 7:275—320. 

Brou, V. A. 1980. New status for Eumorpha intermedia (Sphingidae). J. Lepid. Soc. 
34:302—306. 

. 1992a. Extended duty bait trap designed for continual year-round use. Southern 

Lepid. News 14:4-6. 

. 1992b. Plain talk for entomologists about ultraviolet light. Southern Lepid. News 

14:20-23. 

. 1994. New species of Lapara (Sphingidae) from southeastern United States. J. 
Lepid. Soc. 48:51—57. 

CLARK, B. P. 1917. New Sphingidae. Proc. New England Zool. Cl. 6:57—72 . 

COVELL, C. V., JR. 1984. A field guide to moths of eastern North America. Houghton Mif- 
flin Co., Boston. 469 pp. 

DRAUDT, M. 1931. Family Sphingidae. In A. Seitz (Ed.), The macrolepidoptera of the 
world, Vol. 6. Alfred Kernan, Stuttgart. 

FORBES, W. T. M. 1948. Lepidoptera of New York and neighboring states, Part IT. Cornell 
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FREEMAN, A. 1938. Notes on the Sphingidae (Lepidoptera) of Arkansas. Field and Labo- 
ratory 6:33—43. 

HEITZMAN, J. R. & J. E. HEITZMAN. 1987. Butterflies and moths of Missouri. Missouri 
Dept. Conservation. 385 pp. 

HINE, J. S. 1906. A second contribution on the entomology of the region of the Gulf Bio- 
logic Station. Gulf Biol. Sta. 6:65—83. 

HopcEs, R. W. 1971. The moths of America north of Mexico, Fasc. 21, Sphingoidea. 
E. W. Classey Ltd. & R. B. D. Publications. 158 pp. 


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(Ed). 1983. Check list of the Lepidoptera of America north of Mexico. E. W. Classey 
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HOLLAND, W. J. 1903. The moth book. Doubleday, Page & Co., New York. 479 pp. 

JUNG, R. C. 1950. An annotated list of the Lepidoptera of the New Orleans area. Proc. 
Louisiana Acad. Sci. 8:42—48. 

KOEBELE, A. 1881. Descriptions of and notes upon various larvae. Bull. Brooklyn Ento- 
mol. Soc. 4:20—22. 

LuTz, F. E. 1948. Field book of insects. G. P. Putnam & Sons, New York. 510 pp. 

MERKL, M. E. & T. R. PFRIMMER. 1955. Light trap investigations at Stoneville, Miss., and 
Tallulah, La., during 1954. J. Econ. Entomol. 48:740—741. 

Neck, R. W. 1991. Hawkmoths (Sphingidae) in the Whitley collection from Walker 
County Texas. J. Lepid. Soc. 45:231—233. 

OTTOLENGUI, R. 1894. Entomol. News 5:314. 

PFRIMMER, T. R. 1957. Response of insects to different sources of blacklight. J. Econ. En- 
tomol. 50:801—803. 

PuaTT, A. P. 1969. A lightweight collapsible bait trap for Lepidoptera. J. Lepid. Soc. 
23:97-101. 

RIOTTE, J. C. E. 1972. A review of the North American hawk moth genus Lapara (Lepi- 
doptera: Sphingidae). Life Sci. Contr., Royal Ontario Mus. No. 79:1—40. 

. 1980. Sphinx poecila, a valid North American hawkmoth species (Lepidoptera: 
Sphingidae). Great Lakes Entomol. 13:115—130. 

ROTHSCHILD, W. & K. JORDAN. 1903. A revision of the lepidopterous family Sphingidae. 
Novit. Zool., ix suppl. cxxxv + 972 pp. 

ROWLEY, R. R. 1898. Notes on Missouri sphinges. Entomol. News 9:189—191. 

. 1899. Notes on Missouri Sphingidae. Entomol. News 10:10—12. 

SELMAN, C. L. & H. E. BARTON. 1971. The relative abundance, seasonal distribution and 
taxonomy of the Sphingidae of northeast Arkansas. Ark. Acad. Sci. Proc. 25:56—68. 

TAYLOR, R. & B. TAYLOR. 1965. Collecting sphingids and other moths on the Mississippi 
Gulf Coast. J. Lepid. Soc. 19:189—190. 

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. 1881. Scribner’s Monthly 22:864. 


Received for publication 10 March 1993; revised and accepted 24 March 1996. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
51(2), 1997, 176-179 


CALLOPHRYS ERYPHON (LYCAENIDAE) COLONIZES URBAN AND 
SUBURBAN SAN FRANCISCO BAY AREA, CALIFORNIA, USING 
PLANTED MONTEREY PINE 


Additional key words: Incisalia, biogeography, dispersal. 


The western banded elfin, Callophrys (Incisalia) eryphon (Boisduval), is widespread in 
western North America, mainly in Transition Life Zone and montane regions, where its 
larvae feed on various conifers, primarily Pinaceae (Hardy 1959, McGugan 1958, New- 
comer 1973). In California, this butterfly occurs from the Cascade Range southward along 
both sides of the Sierra Nevada and in the North Coast Ranges, mostly at elevations of 
1000—2500 m, to the San Bernardino and San Jacinto Mountains of southern California 
above 2000 m (Essig Museum specimens). Along the north coast, natural populations of 
C. eryphon range nearly to sea level, near Plantation, Sonoma Co. and Invemess, Marin 
Co., in association with Bishop pine, Pinus muricata. The elfin may have been native on 
the Peninsula south of San Francisco because there are three specimens in the Museum 
of Comparative Zoology, Harvard University, labelled “San Mateo, Cal. A. Agassiz,” prob- 
ably dating from the 19th century. However, there are no modern records from the Penin- 
sula or Santa Cruz Mountains area (Steiner 1990). In Monterey County, a population oc- 
curs at the S. F. B. Morse Botanical Reserve on the Monterey Peninsula in association 
with an isolated colony of native Bishop pine (J. Lane pers. comm., LACM specimens), 
but C. eryphon is not known from Monterey pine (Pinus radiata) there or at the other na- 
tive stands, nor from other native pines of central coastal California. 

There are old records (1929 to 1950) from San Francisco (Steiner 1990); included are 
specimens collected at The Presidio, where Monterey pine has been grown for more than 
a century. However, H. Reinhard (unpubl. data), J. E. Hafernik (in litt.), and I have failed 
to find C. eryphon there in recent years. According to H. H. Behr, conifers grew on Lone 
Mountain at the western edge of the city in the late 1800’s (Howell et al. 1958). These 
likely were Pinus muricata or P. radiata, Howell et al. reasoned, so it is possible that a 
colony of C. eryphon existed there, and its descendants adopted plantings of Monterey 
pine at The Presidio. However, because there are no specimens taken by H. E. Cottle, F. 
X. Williams, or other early 20th century collectors in San Francisco (Steiner 1990), it 
seems likely that the later records represent an adventive colony originating from native 
conifers of Marin County 15-30 km to the northwest, the direction of prevailing winds. 

In recent years the western banded elfin has expanded its range in the San Francisco 
Bay region. It evidently occurred naturally inland in Marin County in association with 
Bishop Pine or Douglas-fir (Pseudotsuga menziesii), because the butterfly was collected at 
Mill Valley on the east side of Mt. Tamalpais in 1908. In recent decades, C. eryphon has 
been discovered successively southeastward from Mt. Tamalpais, in suburban areas dis- 
tant from native conifers: on the eastern bay shore of Marin County at Strawberry Point in 
the grounds of a seminary in 1973 and in an urban yard in Belvedere in 1980; and across 
the bay, at Pt. Molate, Contra Costa Co. in 1989, in association with young Monterey pine 
in a park that was developed in the 1960's. In 1994 and 1995 C. eryphon appeared at sev- 
eral sites on both sides of the Berkeley Hills (Fig. 1). There is no record of this butterfly in 
the East Bay area (Contra Costa and Alameda counties) prior to 1989 (Opler & Langston 
1968, Steiner 1990). 

During 1994, single females were observed in urban gardens in Kensington and Berke- 
ley, and east of the Berkeley Hills at San Pablo Reservoir males perched on understory 
shrubs in a mature Monterey Pine woods planted more than 50 years ago. Additional indi- 
viduals were encountered on four dates in 1995: near Pt. Richmond, on the University of 
California (UC) campus, in Strawberry Canyon at the UC Botanic Garden, and at 425 m 
elevation in the Berkeley Hills near the southern end of Grizzly Peak Blvd. At each of 
these East Bay sites, adults were active in the vicinity of Pinus radiata. 

Any of these populations could have much older origins than the records document. 


VOLUME 51, NUMBER 2 a 


SIERRA 


e 
leveds NEVADA 
ty 


PLACER 


‘> 
iy 


Inverness 
Ridge 


Mt. 
Tamalpais 


CONTRA 
COSTA 


1989 


@ Presumed native 
@ Presumed adventive 


O Negative search 1995 
\ 


O) 


ics VO 
1929-50 


SAN 
FRANCISCO 


Fic. 1. Central California, showing positions of counties mentioned in the text. Inset 
(below) depicts spatial and temporal distribution of Callophrys eryphon in Marin, San 
Francisco, Contra Costa, and Alameda counties adjacent to San Francisco Bay. Presumed 
native colonies in Marin County (filled symbols) are associated with native conifers; dated 
localities (half-filled symbols) refer to adventive colonies associated with Monterey pine 
plantings; no C. eryphon were seen at several peripheral East Bay sites in March—April 
1995 (open symbols). 


Nevertheless, because there is a long history of extensive Monterey pine planting in the 
cities around San Francisco Bay, in gardens, parks, at reservoirs, etc., the lack of older 
records suggests that the recent collections of C. eryphon reflect recent range extension. 
In 1994—95, the butterfly was encountered in the Berkeley area by four observers inde- 
pendently, one a novice collector in an introductory entomology course, each unaware of 
any prior occurrence of C. eryphon in the area. The evidence indicates the elfin has be- 
come established in Contra Costa and Alameda counties in quite recent years. 

If this is true, why did the butterfly fail to colonize Pinus radiata, other than in San 
Francisco, at a much earlier date? Although Callophrys eryphon, Pinus radiata, and P. 
ponderosa are native species in central California, none occurred naturally in the East Bay 
area. Thus, this adaptation to non-native situations may be comparable to that of many in- 


178 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


troduced insects, which undergo a sequence of introduction-establishment, then a long 
period of naturalization, followed by rapid range extension. Such delayed ecogeographical 
expansions are believed to involve increased genetic fitness to environmental conditions to 
which the founder populations were not adapted (e.g., Powell 1983, 1992). Presuming the 
eastward colonization of C. eryphon is recent, it seems reasonable to suppose that this 
handsome butterfly is becoming a widespread urban resident of the East Bay. 

In the Canadian Forest Insect Survey, host tree preferences of C. eryphon in Alberta 
and British Columbia, based on 187 larval collections, were 83% on several species of 
pines (70% on lodgepole pine), 3% on other Pinaceae, namely Douglas-fir and western 
hemlock (Tsuga heterophylla), and 14% on Thuja plicata (Cupressaceae) (McGugan 
1958). In California, there are no records of larval C. eryphon collections from Pinaceae 
other than Pinus (Garth & Tilden 1986, Powell & De Benedictis 1995). 

Collection data for San Francisco Bay region (sr = sight record; sw = slightly worm; w = 
worn): Napa Co.: 2 mi. N Angwin, IV-26-73, IV-17-77, assoc. Pinus ponderosa (R. L. 
Langston). Sonoma Co.: 3 mi. W Plantation, V-5-55 (Langston); 4 mi. W Plantation, V-25- 
57 (J. Powell); Plantation, 800 ft. elev. V-16-58 (O. E. Sette), V-29-60 [P. A. Opler]. Marin 
Co: Inverness, V-18-63 (C. A. Toschi), Inverness Ridge 800—1040', V-15-70, V-10-74, as- 
soc. Pinus muricata (Powell), IV-25-76 (E. Schlinger, M. Helena), IV-22-78 (Powell), IV- 
26-96, in 1995 fire zone (Powell); Mt. Vision, IV-24-82 (Powell); 1 mi. SW Lagunitas, III- 
21-70 (Opler); Mill Valley, IV-4-1908 (F. X. Williams); Strawberry, Golden Gate Baptist 
Seminary, III-28/31-73 (dd 92) (V. & L. Donahue); Belvedere, IV-6-80 (sw 3) (Powell). 
Contra Costa Co.: Pt. Molate Beach, I[V-6-89 (sw 3) (Powell); Pt. Richmond III-16-95 (sw 
°) (Powell); Kensington, IV-13-94 (sr) (Langston); San Pablo Reservoir, IV-18-94 (sw dé) 
(Powell); Berkeley Hills, nr. jct. Grizzly Peak-Skyline Blvds., IV-14-95 (sw 2) (Powell). 
Alameda Co.: Berkeley, nr. La Loma Park, IV-30-94 (w 2) (D. Rubinoff); UC Botanic Gar- 
den, Strawberry Cyn., III-28-95 (sr), IV-10-95 (sw 2), III-6-96 (sr) (Powell); UC Campus, 
IV-5-95 (w 2) (K. Wong). San Francisco Co.: San Francisco, III-29-1929 (W. D. Field), III- 
15-1931 (R. G. Wind), V-5-35 (M. Doudoroff); Presidio, [V-16-49 (L. I. Hewes), IV-12-50 
(E. S. Ross), Presidio nr. Baker Beach, IV-15-50 (J.W. Tilden). 

I appreciate the comments and use of unpublished records provided by: J. P. Donahue, 
Los Angeles County Museum of Natural History (LACM); J. E. Hafernik, California State 
Univ., San Francisco; J. Lane, Santa Cruz, Calif.; R. L. Langston, Kensington, Calif.; P. 
Perkins, Museum of Comparative Zoology, Harvard (MCZ); D. Rubinoff, Univ. Calif. 
Berkeley; and J. Steiner, U. S. Fish and Wildlife Service, Newark, Calif., who compiled 
records from several major collections during his Master's research on San Francisco Bay 
area butterflies. The following reviewed the manuscript and made useful suggestions: P. 
A. Opler, National Biological Service, Ft. Collins, Colo.; R. Robbins, Smithsonian Inst., 
Washington, D.C.; J. Scott, Lakewood, Colo.; and F. A. H. Sperling, Univ. Calif. Berkeley. 


LITERATURE CITED 


GakTH, J. & J. W. TILDEN. 1986. California butterflies. Calif. Nat. Hist. Guides, No. 51. 
Univ. Calif. Press, Berkeley. 246 pp. 

Harpy, G. A. 1959. On the life history of Incisalia eryphon (Lycaenidae) on southern Van- 
couver Island. J. Lepid. Soc. 13:70. 

HowELL, J. T., P. H. RAVEN & P. RUBTZOFF. 1958. A flora of San Francisco, California. 
Wasmann J. Biol. 16:1—157. 

McGucan, B. M. 1958. Forest Lepidoptera of Canada recorded by the Forest Insect Sur- 
vey. Can. Dept. Agric., Forest Biol. Div. Publ. 1034:1—76. 

NEWCOMER, E. J. 1973. Notes on life histories and habits of some western Theclinae. J. 
Lepid. Soc. 27:13-15. 

OPLER, P. A. & R. L. LANGSTON. 1968. A distributional analysis of the gehen ss of Con- 
tra Costa county, California. J. Lepid. Soc. 22:89—107. 

POWELL, J. A. 1983. Expanding geographical and ecological range of Blenageare stultana in 
California (Lepidoptera: Tortricidae). Pan-Pacific Entomol. 59:233—239. 

. 1992. Recent colonization of the San Francisco Bay area, California, by exotic 

imethe (Lepidoptera: Tineoidea, Gelechioidea, Tortricoidea, Pyraloidea). Pan-Pacific 

Entomol. 68:105—121. 


VOLUME 51, NUMBER 2 179 


POWELL, J. A. & J. A. DE BENEDICTIS. 1995. Foliage feeding Lepidoptera of Abies and 
Pseudotsuga associated with Choristoneura in California, pp. 168-215. In Powell, 
J. A. (ed.), Biosystematic studies of conifer-feeding Choristoneura (Lepidoptera: Tor- 
tricidae) in the western United States. Univ. Calif. Publ. Entomol. 115; 275 pp. 

STEINER. 1990. Butterflies of the San Francisco Bay Area. Unpubl. M.S. Thesis, Calif. 
State Univ., Hayward. 93 pp. 


JERRY A. POWELL, Essig Museum of Entomology, University of California, Berkeley, 
California 94720, USA. 


Received for publication 1 November 1995; revised and accepted 13 May 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 179-184 


DIURNAL LEPIDOPTERA OF NATIVE AND 
RECONSTRUCTED PRAIRIES IN EASTERN MINNESOTA 


Additional key words: surveys, species richness, vagility. 


Prairie butterflies are subjects of increasing conservation concern. Their habitat has 
been greatly diminished, and their ability to survive on managed sites and to colonize new 
sites or recolonize old ones is in doubt (Opler 1991). In this paper I report on and com- 
pare the diurnal Lepidoptera communities of both native and reconstructed prairies in 
Minnesota. 

_ I collected insects from the flowers of 58 forb species in four native prairie sites and 
four prairie reconstructions (former agricultural areas recently replanted to prairie) dur- 
ing the summers of 1990, 1991 and 1992. The sites are described in Table 1. Insects were 
collected between 0900 h and 1600 h on sunny or partly cloudy days when the tempera- 
ture was between 20° and 35° C. Collections were made from late May to late September. 
I made one 15 min aerial net collection of insects on the flowers of each forb species with 
at least 100 flowers or inflorescences open, for a total of 507 collections from all forb spe- 
cies in all sites over the three summers. Thus, the number of collections made from a site 
was closely related to the number of forb species present in populations large enough to 
produce 100 or more flowers. Although only a small fraction of the Lepidoptera present 
on a site can be sampled by daylight collections, many of the species of conservation con- 
cern are diurnal. 

The 507 collections yielded 3702 insects representing 305 species; 295 of these were 
identified at least to genus (Reed 1995). There were 118 Lepidoptera individuals repre- 
senting 28 species: 24 butterflies and four diurnal moths (Table 2). Insect vouchers are de- 
posited in the University of Minnesota Insect Museum, and plant vouchers are in the Uni- 
versity of Minnesota Herbarium. 

Collections in native sites produced greater species richness than in reconstructed sites: 
73 individuals and 21 species in 218 15-min collections from native sites, compared to 45 
individuals and 16 species in 289 collections from reconstructions. Five of the 28 species 
collected were described as prairie obligates by Orwig (1992): Callophrys gryneus (Hub- 
ner), Hesperia |. leonardus Harris, H. 1. pawnee Dodge, Polites origines (Fabr.) and 
Satyrium edwardsii (Grote & Robinson) and an additional four species were described as 
remnant-restricted by Panzer et al. (1995): Euphyes conspicua (Edw.) Harkenclenus titus 
(Fabr.), Speyeria aphrodite (Fabr.) and Thorybes pylades (Scudder). Of these nine spe- 
cies, eight were collected from native sites only, none from reconstructions only, and one 
was collected from both native and reconstructed sites. Of the 19 species not considered 
site-restricted, four were collected from native sites only, seven from reconstructions only, 
and eight from both native and reconstructed sites (Table 3). 

Management practices do not appear to account for the differences in species presence 
among sites. There are no obvious differences in management between native sites and 
reconstructions as a group: the large sites are burned in sections, while the small sites 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


180 


a ee EE eS eS oe 


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181 


VOLUME 51, NUMBER 2 


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182 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 2. Scientific and common names of Lepidoptera collected in this study. 
Nomenclature follows Scott (1984), Opler & Krizek (1984) and Covell (1984) 


Hesperiidae 


Atrytone logan (Edw.), Delaware Skipper 

Euphyes conspicua (Edw.), Black Dash 

Euphyes vestris (Boisd.), Dun Skipper 

Hesperia leonardus leonardus Harris, Leonard’s Skipper 
Hesperia leonardus pawnee Dodge, Pawnee Skipper 
Polites coras (Cr.), Peck’s Skipper 

Polites origines (Fabr.), Crossline Skipper 

Polites themistocles (Latr.), Tawny-Edged Skipper 
Wallengrenia egeremet (Scudder), Broken Dash 
Epargyreus clarus (Cr.), Silver Spotted Skipper 
Thorybes pylades (Scudder), Northern Cloudy Wing 


Pieridae 


Colias eurytheme Boisd., Orange Sulphur 
Colias philodice Godart, Clouded Sulphur 


Lycaenidae 


Celastrina ladon (L.), Spring Azure 

Satyrium edwardsii (Gr. & Rob.), Edwards’ Hairstreak 
Callophrys gryneus (Hubner), Olive Hairstreak 
Harkenclenus titus (Fabr.), Coral Hairstreak 


Nymphalidae 


Phyciodes tharos (Drury), Pearl Crescent 

Nymphalis milberti (Godart), Milbert’s Tortoiseshell 
Vanessa cardui (L.), Painted Lady 

Speyeria aphrodite (Fabr.), Aphrodite Fritillary 
Speyeria cybele (Fabr.), Great Spangled Fritillary 
Cercyonis pegala (Fabr.), Wood Nymph 

Asterocampa celtis (Boisd. & Lec.), Hackberry Butterfly 


Sphingidae 


Hemaris diffinis (Boisd.), Snowberry Clearwing 
Hemaris thysbe (Fabr.), Hummingbird Clearwing 


Noctuidae 

Alypia octomaculata Fabr., Eight-Spotted Forester 
Ctenuchidae 

Cisseps fulvicollis (Hubner), Yellow-Collared Scape Moth 


(AREM, CEM, ASP and LLRP) are burned all at once. The ASP and CARP reconstruc- 
tions were mowed for two years following planting, but now are managed by burning. 
Brush cutting is done as needed but does not replace burning on any site. 

It is possible that the reconstructed sites do not provide suitable habitat for these obli- 
gate species. The reconstructions tend to be more mesic than the most species-rich native 
sites (CC and AREM), and five of the eight prairie obligates are reported to be restricted 
to xeric sites by Panzer et al. (1995): Polites origines and Hesperia I. leonardus to xeric 
prairie; Harkenclenus titus to xeric/mesic prairie; Satyrium edwardsii to savanna; and Tho- 
rybes pylades to sand savanna. Hesperia leonardus pawnee and Callophrys gryneus also 
are found in xeric areas (Orwig 1992). Only two of the obligate species collected are re- 
ported by Panzer et al. from mesic sites: Euphyes conspicua from sedge meadow and 


VOLUME 51, NUMBER 2 183 


TABLE 3. Number of individual Lepidoptera species on each prairie site, and their 
nectar plants. Numeric plant abbreviations follow those given in Table 1. Superscript 1 = 
restricted to prairie habitats (Orwig 1992). Superscript 2 = high or moderate remnant re- 
liance (Panzer et al. 1995). 


Native sites Reconstructions 
Species AREM CC CEM LV ASP CARP CHR LLRP Nectar Plants 

Alypia octomaculata 1 34 

Atrytone logan 4 1 Z 3 2 15, 36, 39, 46, 55 

Asterocampa celtis 1 36 

Callophrys gryneus 2 1 42 

Celastrina ladon i 1 

Cercyonis pegala 1 1 1 

Cisseps fulvicollis ® ] 6 1 4 2 1 19) 28) 49AG: 
47, 48, 49, 50 

Colias eurytheme 1 il 1 1 5 Oy he BS BO): 
33, 46, 48 

Colias philodice 1 5 1 91025528 

Epargyreus clarus y) 1 136 

Euphyes conspicua 1 39 

Euphyes vestris 1 5 il 1536442. 46 

Harkenclenus titus? 4 28, 42 

Hemaris diffinis 1 1 3 136555 

Hemaris thysbe 1 il 36 

Hesperia l. leonardus}? 2 28, 31 

Hesperia leonardus pawnee! 2 16 

Nymphalis milberti it 8 

Phyciodes tharos il i 

Polites coras 1 38 

Polites origines}? , il 36 

Polites themistocles i 36 

Satyrium edwardsii\* Aare ell 1, 4, 42, 46 

Speyeria aphrodite? 2 1 1 28, 36 

Speyeria cybele 1 36 

Thorybes pylades? I 1 BO 

Vanessa cardui 3 4 i el TAS 7 2S8e 50: 
58), 

Wallengrenia egeremet gs Teer 1 36 


Speyeria aphrodite from mesic prairie (S. aphrodite was collected from the mesic recon- 
struction CARP—the only obligate individual found on a reconstruction). Beyond these 
associations with general prairie types, specific interactions with foodplants (both larval and 
adult), or larval-tending ants may be required for establishment of certain species, as has 
been demonstrated for other rare Lepidoptera species (Arnold 1983, Cushman & Murphy 
1993). Callophrys gryneus may be absent from the reconstructions due to the absence of 
its larval foodplant, eastern red cedar (Juniperus virginiana) (Opler & Krizek 1984). 

Alternatively, the obligate species may not have reached these reconstructions yet. But- 
terfly populations in some fragmented habitats have diminished mobility (Dempster 
1991), and Cushman and Murphy (1993) suggest that dispersal ability is especially limited 
among lycaenids. Mobility may be influenced by species-specific behavior, such as reluc- 
tance to leave larval foodplants (Arnold 1983). Colonization of new habitat patches by 
these Lepidoptera may be an infrequent event that occurs during “rare years of explosive 
dispersal” as described by Ehrlich and Murphy (1987) for Euphydryas editha. More study 
of the basic biology and mobility of each species is required before we can predict whether 
prairie obligate butterflies will be able to colonize prairie reconstructions. 


184 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


This study was funded in part by a grant from the Non-game Wildlife Division of the 
Minnesota Department of Natural Resources. Field assistance was funded by a Research 
Explorations for Teachers grant to the University of Minnesota Curriculum and Instruc- 
tion Department. I thank Dave Andow, Theresa Leahy, Bill Miller, Susan Weller and the 
site managers for all their help. I especially appreciate the thorough reading and profes- 
sional attitudes of the reviewers. 


LITERATURE CITED 


ARNOLD, R. A. 1983. Ecological studies of six endangered butterflies (Lepidoptera, Ly- 
caenidae): island biogeography, patch dynamics, and the design of habitat preserves. 
Univ. Calif. Publ. Entomol. 99:1—161. 

COVELL, C. V., JR. 1984. A field guide to the moths of eastern North America. Houghton 
Mifflin, Boston. 496 pp. 

CUSHMAN, J. H. & D. D. Murpny. 1993. Susceptibility of lycaenid butterflies to endan- 
germent. Wings (Xerces Society) 17:16—21. 

DEMPSTER, J. P. 1991. Fragmentation, isolation and mobility of insect populations, 
pp. 143-154. In Collins, N. M. & J. A. Thomas (Eds.), The conservation of insects and 
their habitats. Academic Press, London. 

EHRLICH, P. R. & D. D. Murpny. 1987. Conservation lessons from long term studies of 
checkerspot butterflies. Cons. Biol. 1:122—131. 

GREAT PLAINS FLORA ASSOCIATION. 1986. Flora of the Great Plains. Univ. Kansas Press, 
Lawrence, Kansas. 1402 pp. 

OPLER, P. A. 1991. North American problems and perspectives in insect conservation, pp. 
9-32. In Collins, N. M. & J. A. Thomas (Eds.), The conservation of insects and their 
habitats. Academic Press, London. 

OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins 
University Press, Baltimore. 294 pp: 

Orwic, T. T. 1992. Loess hills prairies and butterfly survivia: opportunities and challenges, 
pp. 132-135. In Smith, D. D. & C. A. Jacobs (Eds.), Proceedings of the Twelfth 
North American Prairie Conference. Univ. Northern Iowa, Cedar Falls, Iowa. 

PANZER, R., D. STILLWAUGH, R. GNAEDINGER & G. DERKOVITZ. 1995. Prevalence of rem- 
nant-reliance among the prairie and savanna-inhabiting insects of the Chicago region. 
Natural Areas Journal 15:101—116. 

REED, C. C. 1995. Insects surveyed on flowers in native and reconstructed prairies (Min- 
nesota). Restoration and Management Notes 13:210—213. 

ScoTT, J. A. 1986. The butterflies of North America: a natural history and field guide. 
Stanford Univ. Press, Stanford, California. 585 pp. 


CATHERINE C. REED, Entomology Department, 219 Hodson Hall, University of Min- 
nesota, Saint Paul, Minnesota 55108, USA. 


Received for publication 20 December 1994; revised and accepted 12 March 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 184-187 


YOU CAUGHT WHAT IN YOUR BACKYARD? 


Additional key words: Electrostrymon angelia, Ministrymon azia, Dryas iulia, 
Florida, dispersal. 


What butterflies are in your back yard? This question has been asked before in the 
pages of the Journal (Howe 1959) and many subsequent notes. Howe identified 64 butter- 
fly species on a nine-acre plot in Kansas, at the time a truly impressive feat. We also 


VOLUME 51, NUMBER 2 185 


Fic. 1. Electrostrymon angelia angelia. Male, upper (left) and under (right) surfaces. 
Florida: Manatee Co.; 2 mi. E of Samoset, March 1992 (leg. J. Y. Miller). 


been informally monitoring the butterfly faunas in two localities adjacent or close to our 
respective homes on the Florida Gulf Coast, one in Manatee County (Millers) and the 
other in Pinellas County (Anderson). These studies are recreational rather than scientific 
and undertaken more for curiosity than for any other reason. 

The butterfly fauna of each area was reasonably well known as of early 1992, so it was 
surprising that each of us independently collected, during the latter half ‘of that year, two 
species previously unrecorded from the west coastal area of the state. Both of these were 
lyeaenids, and both have been recorded only recently from southeastern Florida and the 
Florida Keys. 

Electrostrymon angelia angelia (Hewitson) has become a recent resident in southeast- 
erm Florida (Anderson 1974), and its known range extends along the Atlantic coast to 
about Fort Lauderdale. In March, and again in October 1992, Lee and Jackie Miller en- 
countered this small hairstreak in their backyard nectaring on flowers of Hyptis verticil- 
lata Jacq. (Lamiaceae) (determination by R. Wunderlin, Univ. So. Florida). Anderson first 
found this species in Pinellas County in November 1992, nectaring on flowers of golden- 
rod, Solidago sp. (Asteraceae). Since the first sightings of this butterfly on the west coast 
of Florida were in March 1992, prior to the destruction wrought by Hurricane Andrew in 
southern Florida in August 1992, that storm could not have been the cause of this range 
extension. These specimens, one of which is illustrated here (Fig. 1), represent the first 
records of E. angelia for Pinellas and Manatee Counties, and resident populations, al- 
though quite small, have been seen in the area until present. The species was previously 
reported in Lee County on the Florida Gulf Coast by Heinrich in 1989 (see Baggett 1989). 

Specimens of Ministrymon azia (Hewitson) were collected by Anderson during No- 
vember 1992 in Pinellas County (Figs. 2, 3). About the same time in Manatee County, Lee 
and Jackie Miller made two positive sight records, with an additional sight record on the 
grounds of the Allyn Museum in Sarasota, Sarasota County. The first Manatee County 
specimen, a female, flew out of a tree and landed on the windshield of a car during a cool 
day; then it proceeded to thermoregulate there for several minutes with its wings alter- 
nately opening and closing, thus revealing the diagnostic ventral red spotband, and the 
gray-powdered upper hindwing that established its sex. These specimens, or their ances- 
tors, might have been introduced through the actions of Hurricane Andrew, but due to 
their fresh condition, we suspect the species has been resident longer and simply avoided 
detection because of its small size and similar appearance to Leptotes cassius theonus (Lu- 
cas). There also is one record of M. azia from New Port Richey, Pasco County (Baggett 
1989) captured late that year. 

Both E. angelia and M. azia will feed as larvae on Brazilian pepper, Schinus terebinthi- 
folius Raddi (Anacardiaceae), a ubiquitous weed in southern peninsular Florida that is well 


186 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 2-3. Ministrymon azia. 2, male; 3, female (3). Upper (left) and under (right) 
surfaces. Both Florida: Pinellas Co.; St. Petersburg, November 1992 (leg. R. A. Anderson). 


established on the Miller property, and M. azia also is known to feed upon Leucaena leu- 
cocephala (Lam.) de Wit (Fabaceae). It seems likely that there has been an established 
breeding population of one or both species for several years during a series of consecutive 
extremely mild winters since 1989. Because both butterflies have broad ranges and dis- 
perse readily throughout the Caribbean (Smith et al. 1994) and have recently become es- 
tablished in Florida, it is likely that these butterflies arrived in west coastal Florida by nat- 
ural dispersal. Lee and Jackie Miller have seen E. angelia every year since, through the 
spring of 1996, and Anderson has taken both species in Pinellas County, so apparently 
both species are still firmly established. The butterflies certainly are not limited by their 
anacardiaceous foodplant, which ranges to near Clearwater in northern Pinellas County, 
and both species should be sought elsewhere in southwestern Florida. 

Lee and Jackie Miller also observed a specimen of Dryas iulia largo Clench on 28 Feb- 
ruary taking nectar at Citrus flowers. It was captured, found to be a ragged male, and un- 
fortunately released before the real significance of the record was realized, as D. i. largo 
was known previously only from extreme southern Florida (Kimball 1965). The Manatee 
County specimen was observed farther north in west coastal Florida than any previous 
record, although a recent sighting in Orlando by Deuerling (see Baggett 1993) would sug- 
gest that this is another species actively expanding its range, possibly during the recent 
warm winters. 

If there is a moral to be learned from this tale, it is that one can never say with confi- 
dence that one knows everything about the distribution of butterflies in an area. Many 
species may expand their ranges when conditions are favorable only to have the ranges 
contract subsequently. It will be intriguing to see whether the populations noted here can 
persist after a cooler winter with several frosts. There are many examples in the literature 
of transient populations of animals from many parts of the world. Armadillos and oppos- 
sums are well-known examples of such prior dispersals in North America, and we have ob- 


VOLUME 51, NUMBER 2 187 


served expansions and contractions of butterflies such as Calpodes ethlius (Stoll) and 
Siproeta stelenes biplagiata (Fruhstorfer) in Central Florida. Because of larval hostplant 
relations, we also must remember that larvae might inadvertantly be transported with 
nursery plants from different areas within the state. However, the lycaenid records listed 
here were made long after the active growth period when exotic plants would normally be 
brought into Central Florida for sale in local nurseries. 

We consider voucher specimens to be an absolute necessity in faunal survey studies in 
order to adequately determine the taxa represented in an area. Vouchers of M. azia and E. 
angelia discussed here have been deposited in the collections of the Allyn Museum of En- 
tomology, Florida Museum of Natural History. 


LITERATURE CITED 


ANDERSON, R. A. 1974. Three new United States records (Lycaenidae and Nymphalidae) 
and other unusual captures from the lower Florida Keys. J. Lepid. Soc. 28:354—358. 

BAGGETT, H. D. 1989. Current Zone Reports. So. Lepid. News 11(4):44. 

. 1993. Current Zone Reports. So. Lepid. News 15(2):21. 

HoweE, W. H. (ed.) 1958. What’s in your backyard? Lepid. News 12:130. 

KIMBALL, C. P. 1965. The Lepidoptera of Florida. An annotated checklist. State of Florida 
Dept. Agric., Gainesville, Florida. 353 pp. 

SMITH, D. S., L. D. MILLER & J. Y. MILLER. 1994. The Butterflies of the West Indies and 
South Florida. Oxford Univ. Press. 264 pp. 


LEE D. MILLER, JACQUELINE Y. MILLER, Allyn Museum of Entomology of the Florida 
Museum of Natural History, 3621 Bay Shore Road, Sarasota, Florida 34234, USA, AND 
RICHARD A. ANDERSON, 836 Amelia Court NE, St. Petersburg, Florida 33702, USA 


Received for publication 1 May 1994; revised and accepted 25 August 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 187-190 


REPRODUCTIVE ADAPTATIONS OF THE TASAR SILKMOTH, ANTHERAEA 
MYLITTA (SATURNIIDAE), TO EMERGENCE SEASON 


Additional key words: ovary, coupling, fecundity, hatching, diapause. 


Most insects survive periods of environmental stress by entering a state of diapause. 
The Indian tropical tasar silkworm, Antheraea mylitta Drury, completes two to three gen- 
erations in a year (Sinha & Chaudhuri 1992), and in bi/trivoltine broods undergoes pupal 
diapause for a period of about six to seven months to overcome unfavorable environmen- 
tal conditions (Dash & Nayak 1988, Kapila et al. 1991, Sinha & Chaudhuri 1992). Pupal 
diapause in this species normally terminates at the end of May and eclosion begins in June 
with the advent of rain (Sinha & Chaudhuri 1992). This is known as optimal seasonal 
emergence. However, in the diapausing brood a portion of the pupae hatch 1—2 months 
early, emerging in a presumably unfavorable climate before the rainy season (Kapila et al. 
1991). The physiological/hormonal basis of this erratic eclosion remains unclear, although 
endocrine regulation of pupal diapause in other insects has been well documented 
(Browning 1981, Denlinger 1985). Daily patterns of insect behavior (e.g., locomotion, 
feeding, emergence, mating, oviposition, and hatching) are governed by daily cycles of 
temperature, humidity, and light intensity as well as by physiological events (Beck 1983, 
Ratte 1985, Ashby & Singh 1990). We report here on ovary morphology and reproductive 
behavior of “seasonally” and “unseasonally” emerged tasar silk moths. 

One thousand diapausing A. mylitta pupae of each sex were observed as they emerged 


188 


TABLE 1. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


and unseasonally emerged moths (P < 0.05, t-tests). 


Emergence and reproductive parameters for seasonally and unseasonally 
emerged Antheraea mylitta. Asterisks indicate significant differences between seasonally 


Parameters Seasonal Unseasonal 
% Emergence 
male 62.80 0.70 
female 67.90 0.90 
total 65.35 0.80 
% Coupling 
self 47.42 Tat 
mechanical 37.70 5b b> 
total 85.12 66.66 
Female moth weight (gm)* 6160.12 + 110.95 4015.50 + 210.97 
Ovary weight (mg)* 3598.75 + 98.08 2092.50 + 55.87 
Single egg weight (mg)* 10.05 + 0.10 8.07 + 0.13 
Number of eggs 
laid* 162 +5 10+1 
unlaid 4444 42 +1 
total* 206 + 8 52 +2 
% Eggs 
laid* 78.96 + 1.59 19.59 + 1.71 
unlaid* 21.04 + 1.59 80.41 + 1.71 
Incubation period (d)* 8.55 + 0.15 7.50 + 0.20 
% Hatching* 80.76 + 1.58 18.48 + 3.93 
Average temperature (°C) 
minimum* 26.80 + 0.31 23.10 + 0.28 
maximum* 30.36 + 0.26 32.97 + 0.24 
Average relative humidity* U3 AS 2 127 39.96 + 1.73 


in 1991. These had fed previously as larvae on the leaves of Terminalia arjuna. Moths that 
emerged during March—April were considered “unseasonal” whereas those that emerged 
during June-July were considered “seasonal.” Emergence and mating percentage were 
recorded for each group. Moths were allowed to pair naturally for about three or four 
hours; adult females unable to mate were coupled manually. Subsequently, the moths 
were allowed to oviposit. Egg production was recorded by counting the number of eggs 
laid by females, and the number of unlaid eggs was determined by dissection. Ovary mor- 
phology was assessed after removing the whole system from the adult moth. Temperature 
and relative humidity were recorded throughout the observation period. 

Table 1 shows variation in reproductive parameters in relation to emergence. Unsea- 
sonal eclosion was rare in comparison to seasonal eclosion, and the natural coupling per- 
centage of unseasonally emerged moths was lower. Unseasonally emerged female moths 
had lower body weight, lower total ovary weight, lower single egg weight, and decreased 
egg production, egg laying capacity and egg hatching. Table 2 indicates that unseasonally 
emerged moths had smaller ovaries, ovarioles, and mature eggs. Unseasonally emerged 


TABLE 2. Physical dimensions of ovary, ovarioles and mature eggs (after laying) of un- 
seasonally and seasonally emerged Antheraea mylitta. Significant differences between sea- 
sonally and unseasonally emerged moths existed for all traits measured (P < 0.05, t-tests). 


Ovary Ovariole Mature eggs 


Length Width Length Width Length Width 


Seasonal 55.67+0.54 14.67+0.27 84.25+058 3.07+0.03 3.044+0.04 2.53 + 0.02 
Unseasonal 42.33 +1.19 11.33+0.54 65.12+061 250+0.02 254+0.04 2.19 + 0.04 


VOLUME 51, NUMBER 2 189 


Fic. 1. Ovaries of seasonally (left) and unseasonally (right) emerged Antheraea 
mylitta. 


moths also frequently had oocyte-free zones when compared to seasonally emerged moths 
(Fig. 1). 

a fluctuated more during March—April than during the months of June— 
July, and relative humidity was lower in March—April. Variation in ambient environmental 
factors may be responsible for some of the observed differences in reproductive biology in 
A. mylitta (see e.g., Messenger 1964, Hagstrum & Leach 1973, Beck 1983, Sidibe & 
Lauge 1977, Ratte 1985, Ochieng’-Odero 1991), but we do not yet know why pupal dia- 
pause occasionally terminates early in this species. Biogenic amines have been implicated 
in the regulation of development, especially in diapause induction and termination 
(Puiroux et al. 1990), and our own unpublished observations suggest that octopamine plays 
a major role in termination of pupal diapause in A. mylitta. Unseasonally emerging tasar 
silk moths are not being exploited commercially at present. 

We thank the Director of the Central Tasar Research and Training Institute, Ranchi, 
for providing research facilities. 


LITERATURE CITED 


ASHBY, M. D. & P. SINGH. 1990. Control of diapause in codling moth larvae. Entomol. 
Exp. Appl. 56:71—81. 

BECK, S. D. 1980. Insect photoperiodism. 2nd Ed. Academic Press, New York. 160 pp. 

. 1983. Thermal and thermoperiodic effects on larval development and diapause in 
the European corn borer, Ostrinia nubilalis. J. Insect Physiol. 29:107—112. 

BROWNING, T. O. 1981. Ecdysteroids and diapause in pupae of Heliothis punctiger. J. In- 
sect Physiol. 27:715—719. 

DasH, A. K. & B. K. Nayak. 1988. Effect of refrigeration on hatching of eggs of the tasar 
silk moth, Antheraea mylitta Drury (Saturniidae). J. Res. Lepid. 27:263—265. 

DENLINGER, D. L. 1985. Hormonal control of diapause, pp. 353-412. In Kerkut, G. A. & 
L. I. Gilbert (Eds.), Comprehensive insect physiology, biochemistry, and pharmacol- 
ogy. Vol. 8. Pergamon Press, Oxford. 

HacstTruM, D. W. & C. E. LEACH. 1973. Role of constant and fluctuating temperature in 
determining development time and fecundity of three species of stored-product 
Coleoptera. Ann. Entomol. Soc. Am. 66:407—409. 

KAPILA, M. L., A. CHAUDHURI, O. P. DUBEY, C. C. CHAUDHURI & S. S. SINHA. 1991. 
Studies on the preservation of seed cocoons of tasar silkworm, Antheraea mylitta D. 
during diapause. Sericologia 32:579—591. 


190 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


MESSENGER, P. S. 1964. The influence of rhythmically fluctuating temperatures on the de- 
velopment and reproduction of the spotted alfalfa aphid, Themen maculata. J. 
Econ. Entomol. 57:71—76. 

OCHIENG’-ODERO, J. P. R. 1991. The effect of photoperiod and thermophotoperiod on the 
larval critical weight, latent feeding period, larval maximum weight and fecundity of 
Cnephasia jactatana (Walker) (Lepidoptera: Tortricidae). J. Insect. Physiol. 
37:441—445. 

Purrox, J., R. MOREAU & L. GoRDOUx. 1990. Variation of biogenic amine levels in the 
brain of Pieris brassicae pupae during non-diapausing and diapausing development. 
Arch. Insect. Biochem. Physiol. 14:57—69. 

SIDIBE, B. & G. LAUGE. 1977. Incidence de thermoperiodes et de temperatures con- 
stantes sur quelque criteres biologique de Spodoptera littoralis Boisduval (Lepi- 
doptera, Noctuidae). Ann. Soc. ent. France 13:369—379. 

SINHA, A. K. & A. CHAUDHURI. 1992. Factors influencing the phenology of different 
broods of tropical tasar silkmoth Antheraea mylitta Drury (Lepidoptera: Saturniidae) 
in relation to its emergence and post emergence behavior. Environ. Ecol. 
10:952—958. 


ANATHBANDHU CHAUDHURI, Central Sericultural Research and Training Institute, 
Berhampore 742 101, West Bengal, India, DIPANKAR CHAKRABORTY AND ASHOK KUMAR 
SINHA, Central Tasar Research and Training Institute, Ranchi 835 303, Bihar, India. 


Received for publication 1 March 1994; revised and accepted 24 June 1996. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 191-193 


BOOK REVIEWS 


MARIPOSAS DE CHIAPAS, by Roberto G. de la Maza E. and Javier de la Maza E. 1993. 
Gobierno del Estado de Chiapas. Distributed by Montes Azules, Camino Real a Xochi- 
milo No. 60, Tepepan Xochimilo 16020, Mexico DF, Mexico (email: bfly@sar.net). 223 pp., 
152 color plates. Hardcover, dustjacket, 21.5 x 28.5 cm, ISBN 968-6258-34-5. $60 US 
(postpaid). 


Chiapas holds a special place in my heart—my first experience of the neotropics was a 
month-long unstructured tour of the state. I return every few years to recapture the ex- 
citement I felt during that month-long adrenaline rush, and Chiapas never fails me. It is a 
land of contrasts, culturally and biologically the richest state in one of the most diverse 
countries in the world, yet its neglected economy and ecologically abused highlands con- 
tribute to an air of depression. Its habitats run the gamut: from lowland rain forests and 
swamps, through the rich forests of the lower elevation hills, and upwards to the highland 
deserts, oak/pine forests, and tropical cloud forests along mountainous ridges. Its highland 
butterflies include species more familiar to Canadians, such as Nymphalis antiopa, Papilio 
polyxenes, and Colias eurytheme, whereas in the lowlands, classic Neotropical genera 
abound, such as Morpho, Memphis, and Parides. Thus, it is fitting that this marvelous vol- 
ume is defined by its contradictions, characterized in many respects by what it is not rather 
than what it is. It is not an identification guide, but it is a picture book with over 100 
mostly superb photographs of butterflies in their natural habitats. It is written for local 
consumption as a general introduction to butterflies, and yet it contains a storehouse of 
natural history and distribution information that will be used throughout Central America 
and beyond. It is not, however, an easy book from which to extract this valuable informa- 
tion, and most of the species-specific information is presented in tabular format. 

In the tradition of the family’s Mariposas Mexicanas (R. de la Maza R., 1987, Mariposas 
Mexicanas, Guia para su Colecta y Determinacion, 302 pp., 67 color pls.), the current vol- 
ume features not just butterflies, but the context in which they occur, devoting 170 pages 
of text and photographs to historical and ecological context. The book opens with an brief 
overview of butterfly life history and ecology, but quickly focuses on Chiapas ecosystems 
and biogeography. The bulk of this introductory section details the historical context that 
produced this volume—the significant field collectors involved, and the interesting history 
of involvement of the entire de la Maza clan with Chiapas butterflies. Have no doubt 
about it, this book is as comprehensive an effort as has ever been mounted in Central 
America. The authors live in Chiapas, and this volume has been an obvious labor of love 
for many years. The field work and experience that underpins this volume is unequaled 
for Central America. 

The heart of the book is the chapter entitled “La fauna de mariposas chiapanecas,’ 
which includes a powerful trip through the major ecological and faunal butterfly commu- 
nities of Chiapas. First, the authors de-construct faunal communities into those of dis- 
turbed habitats and those of ‘stable ecosystems —a division that literally separates species 
that thrive in highly disrupted landscapes from those that require relatively undisturbed 
habitats. These two categories are each divided further, the disturbed fauna by altitude, 
the habitat-restricted fauna by broad vegetational cover types, which seem to be strongly 
influenced by altitude, rainfall, and biogeographic affinities. Each faunal community is dis- 
cussed relative to the primary factors that influence the habitats found within it, and each 
discussion is illustrated with excellent maps, habitat photographs, and some excellent pho- 
tos of the more interesting butterflies found there, such as those that are the most habitat 
restricted or which are characteristic of the fauna. This section includes many photos of 
species that are rarely seen. This chapter continues with a curious attempt to group but- 
terflies based on broad evaluations of color patterns and which portion of the habitat they 
use. For example, “Patrén banda oblicua’ (which I broadly translate as ‘butterflies with an 
oblique band through their forewing, mostly nymphalids but including some metalmarks 
and skippers) are found primarily in the sub-canopy of forest communities. Although there 
may in fact be some truth to these generalizations, the exceptions drive you mad, and the 


192 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


end result depends to no small degree on difficult decisions of where to place a given but- 
terfly species in these 21 pattern groups. The subjective quality of the groups allows the 
reader considerable latitude to move species around among the groups to fit one’s precon- 
ceptions, although the arrangement is certainly thought-provoking. This chapter ends with 
an overview of butterfly life histories and the conservation status of the Chiapas fauna as 
well as the threats to biodiversity that face the state. 

The final chapter is a beguiling overview of butterfly classification, which opens with a 
comparison of the classification system used in the village of Tzeltal for the conspicuous 
kinds of butterflies (large, small, white, blue, etc.) relative to traditional Linnean classifica- 
tion. The remainder of the chapter is devoted to an overview of the families and subfami- 
lies of butterflies in Chiapas. Although most taxonomists may fret about the classification 
followed in this book, my opinion is that it doesn’t detract from the utility of the volume. 

The appendices are what I find most interesting. Appendix II is a simple listing of 52 
species that are in danger of extinction in Chiapas. The list includes a few species with 
which I am familiar, but most of these species I have never encountered in the state. Be- 
cause this is simply a list of species, it is difficult to determine if these species are truly im- 
periled with identifiable threats, or are simply very rarely encountered. I suspect that the 
list includes both, but generic threats to biodiversity abound in Chiapas, and I have no 
doubt that deforestation of highland and tropical forests threaten many of the locally dis- 
tributed species. 

In my view, Appendix I is the technical core of this book—a 34-page listing of the 1,194 
species known from Chiapas. No listing like this from the Neotropics will ever be 100% 
complete, but certainly this is the best so far published for the region. (There are at least 
two additional skippers from the state that I am aware of, and I am sure other readers will 
find records in their collections, too, although the additions to be made are certainly mi- 
nor.) This appendix is the first complete faunal list, including all butterfly families, from a 
discrete Central American region, so this information is illuminating. 

But Appendix I is more than just a list of species—five columns of data are included for 
each. The first column divides the state into nine zones of distribution, and lists the status 
of each species and subspecies in each zone (several taxa are represented by two sub- 
species in Chiapas, thanks to mountainous terrain that effectively divides the lowland trop- 
ical ecosystems of the state). Status within each of these regions is coded as ‘established, 
present, extinct, dubious, or requires confirmation, thus providing some insight into each 
species’ abundance. Next is a series of codes that tell you to which of the faunal group(s) 
the butterfly belongs. Because the distributions of these faunal groups are mapped, this 
information can be combined with the nine zones of distribution to develop a pretty re- 
fined guess as to the actual (or potential) range of each species within Chiapas. The third 
column’s codes refer to that system of wing-coloration and habitat-use grouping that I 
mentioned previously. I have absolutely no idea what column 4 stands for—it is labeled 
PC and I simply can’t find any information about it or about the codes listed under this 
heading. I am not sure if the explanation got dropped during editing or if my broken Span- 
ish prevents me from finding this information. Either way, this is a vexing problem. The fi- 
nal column presents the attitudinal range from which the species is known (which can be 
added to columns | and 2 to further refine range estimates). In summary, this valuable ap- 
pendix provides a wealth of information and I find myself referring to it on a regular basis. 

I cannot resist the urge to make a simple faunal comparison between Chiapas and 
Costa Rica. In total, Chiapas has fewer species of Nymphalidae, Pieridae, and Papilion- 
idae than does Costa Rica: ca. 450 species compared to Costa Rica’s ca. 550 (P. J. DeVries, 
1987, The Butterflies of Costa Rica and their Natural History: Papilionidae, Pieridae, 
Nymphalidae, Princeton University Press, 327 pp., 50 color pls.). But what this volume 
sharply defines is the magnitude of the butterfly fauna not covered by DeVries—Ly- 
caenidae, Riodinidae, and Hesperiidae, namely, the two-thirds of the butterfly fauna that 
is more difficult to adequately inventory and identify. In this regard, the Chiapas volume 
truly fills a gap in our knowledge of Mesoamerican butterflies and allows us to better pon- 
der the realities of butterfly diversity in the region. 

So, do I recommend this volume? Unequivocally, yes. It has been quite a while since I 


VOLUME 51, NUMBER 2 193 


purchased a book in this price range that did not disappoint—usually such books are all 
fluff and no content. At twice the price, Mariposas de Chiapas would be a bargain. 


OHN A. SHUEY, The Nature Conservancy, 1330 West 38th Street, Indianapolis, Indi- 
ana 46208, USA. 


Journal of the Lepidopterists’ Society 
51(2), 1997, 193-194 


CARCASSON’S AFRICAN BUTTERFLIES: AN ANNOTATED CATALOGUE OF THE PAPIL- 
IONOIDEA AND HESPERIOIDEA OF THE AFROTROPICAL REGION, edited by P. R. Ackery, C. 
R. Smith, and R. I. Vane-Wright. 1995. CSIRO Publications. ix + 803 pp., 300 black-and- 
white photographs. Hardcover, 27.5 x 21.0 cm, ISBN 0-643-05561-4. $150. (In North 
America, order from ISBN, 5602 NE Hassalo Street, Portland, OR 97213-3640.) 


Upon first glimpse of the title, one may ask: “Who is this fellow Carcasson, and which 
are his African butterflies?” The answers to these questions (and many others) are found 
on the first few introductory pages of this handsome book. Bob Carcasson was an English- 
born entomologist who spent a considerable portion of his highly productive career in 
Africa, studying the Lepidoptera of that continent. A draft manuscript prepared by Car- 
casson over 25 years ago was the nucleus upon which this extremely thorough treatment 
was built. The final product is the result of the masterful editorship and knowledge of 
three outstanding British lepidopterists—P. Ackery, C. Smith, and R. Vane-Wright. 

This annotated (and illustrated) catalogue represents the first comprehensive treatment 
of the butterfly fauna (Papilionoidea and Herperioidea) of any large tropical region. Car- 
casson defined the Afrotropical Region (equivalent to the Ethiopian Region of other au- 
thors) on the basis of zoogeography rather than political boundaries; hence, the butterflies 
of North African countries such as Morocco, Libya, and Egypt are not included. The cata- 
logue includes over 3600 species representing a whopping 20% of the world’s butterfly 
fauna. (Among the major faunal realms, the Afrotropical Region supports the third richest 
butterfly fauna, following the Neotropical with approximately 7900 species and the Orien- 
tal with approximately 4100 species.) A representative of each of the 300 genera treated is 
illustrated in a life-sized, black-and-white photograph at the beginning of each generic 
treatment. The catalogue includes all generic, specific, and infraspecific names of the but- 
terflies of the region, organized in a highly usable fashion. Families, subfamilies, and gen- 
era are arranged phylogenetically, with species names arranged alphabetically under each 
genus. There are about 14000 names in the catalogue, including all described species, syn- 
onyms, forms, etc. 

Under each species citation is a reference to the original description and brief notes on 
its distribution. Where known, information on host plants and other biological features is 
summarized. 

The text of the book begins with a brief introduction and a section on general butterfly 
biology in the Afrotropical Region. Next there is an extensive gazetteer, which is particu- 
larly useful given the instability of place names in Africa over the past 50 years. Following 
is a section on biogeography that is an updated and slightly modified version of one of Car- 
casson’s most influential publications—A preliminary survey of the zoogeography of 
African butterflies. The next 620 pages are dedicated to the catalogue itself. The book in- 
cludes a comprehensive index and an up-to-date bibliography on African butterflies. 

There is little to criticize in this beautifully produced work. My only complaint is that 
there seems to be a considerable amount of wasted space. Numerous pages are only half- 
filled with text, and a few have only two or three lines. (This is an extremely trivial com- 
plaint for such a large work.) 

In summary, Carcasson’s African Butterflies is an extremely thorough, well-organized, 
highly usable catalogue of the butterflies of the Afrotropical Region. The introductory ma- 


194 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


terial and illustrations enhance considerably its usefullness and value. The book should 
have a wide appeal to professionals and amateurs interested in any aspect of the butter- 
flies or zoogeography of the Afrotropical Region. : 


JOHN W. Brown, Systematic Entomology Laboratory, ARS, PSI, USDA, °% National 
Museum of Natural History, MRC 168, Washington, DC 20560, USA. 


Journal of the Lepidopterists’ Society 
DCA) 1997, 195 


CORRECTION TO VOLUME 51 


In the General Note by A. K. Sengupta and A. A. Siddiqui, “Effects of gene-environment 
interaction on silk yield in Antheraea mylitta (Saturniidae),” which appeared in 51(1):95— 
97, the captions for both Tables 1 and 2 carry the incorrect taxon. The name Bombyx mori 
appears erroneously instead of Antheraea muylitta. 


Date of Issue (Vol. 51, No. 2): 11 July 1997 


4 
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EDITORIAL STAFF OF THE JOURNAL 


Lawrence F. Gat, Editor 
Computer Systems Office 
Peabody Museum of Natural History 
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Associate Editors: 
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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


THE IMMATURE STAGES OF OXYTENIS MODESTIA, WITH COMMENTS ON THE LAR- 
VAE OF ASTHENIDIA AND Homoeopreryx (SATURNIIDAE: OXYTENINAE) 
Annetie Aiello and Manuel A. Balcazar Lo -.  e 


A NEW SPECIES OF PHANETA, WITH TAXONOMIC DIAGNOSES AND SEASONAL 


AND GEOGRAPHICAL DATA ON FOUR RELATED SPECIES (TORTRICIDAE) 
Donald J. Wright, Richard L. Brown and Loran D. Gibson 


THREE ADDITIONAL BACTRA IN CALIFORNIA, ONE NATIVE BUT OVERLOOKED, 


ONE PROBABLY INTRODUCED, ONE NEW SPECIES (TortricipaE) Jerry A. 
Powell 2 LEA SES IN SOD OTE ace 


A NEW CHAETAGLAEA FROM THE SOUTHEASTERN UnitepD States (NocTulipDae: 
CocunLuNnsr) Vernon Antoine Brou, Jr_. ; 


LarvaL Hosts OF UresipHiTa HUBNER (CRAMBIDAE) Rosemary Leen. 


Host speciricity or UResipHiTA REVERSALIS (GuENEE) (CRAMBIDAE) 
Rosemary Leen. a Oe 


DIsTRIBUTION AND PHENOLOGIES OF LOUISIANA SPHINGIDAE Vernon Antoine 
Brow, [r-and Charlotte Dozar Brow.) 


GENERAL NOTES 


Callophrys eryphon (Lycaenidae) colonizes urban and suburban San Francisco Bay 
area, California, using planted Monterey Pine Jerry A. Powell. 


Diurnal Lepidoptera of native and reconstructed prairies in eastern Minnesota 
Catherine:C: Reed 02 ee OO A a 


You caught what in your backyard? Lee D. Miller, Jacqueline Y. Miller and Richard A. 
Anderson! sito Oo) Wi pa EO ee | a 


Reproductive adaptations of the tasar silkmoth, Antheraea mylitta (Saturniidae), to 
emergence season Anathbandhu Chaudhuri, Dipankar Chakraborty and Ashok 
Kamar Sinha oh gle bo Wa ae ieee 


Book ReEvIEws 


Mariposas de Chiapas: John A;Shuey 2) 0 8 


Carcasson's‘African Butterflies: John Wi Brown) eee 


105 


119 


128 


135 


139 
149 


156 


176 
Whe) 


184 


This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). 


Volume 51 | 1997 Number 3 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 


Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


Sn SONAR 
pre 17.1997» 


5 December 1997 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE CoUNCIL 


James P. Tutte, President CLauDE Lemaire, Vice President 
Eric H. Merzier, Immediate Past Mocens C. NIELsEN, 

President Vice President 
Vitor Osmar BEcKER, Vice President Davip C. IFTNer, Treasurer 


Micuac- J. SMITH, Secretary 


Members at large: 


Richard L. Brown Ronald L. Rutowski M. Deane Bowers 
Charles V. Covell, Jr. Felix A. H. Sperling Ron Leuschner 
John W. Peacock Andrew D. Warren Michael Toliver 


E;DITORIAL BOARD 


Rosert K. Rossins (Chairman), Joun W. Brown (Member at large) 
LawrENcE F. Gatu (Journal) 
WiiuiaM E. Miter (Memoirs) 
Puiu J. ScHappert (News) 


Honorary LirE MEMBERS OF THE SOCIETY 


Cuares L. Reminctron (1966), E. G. Munroe (1973), 
ZpRAVKO Lorxovic (1980), Ian F. B. Common (1987), Jon G. Franctemont (1988) 
Lincoin P. Brower (1990), Douctas C. Fercuson (1990), 
Hon. Miriam Rotuscuitp (1991), Ciraupe Lemaire (1992) 


> 


The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted 
in December 1950, is “to promote the science of lepidopterology in all its branches, . . . to issue a pe- 
riodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by 
both the professional worker and the amateur in the field; to secure cooperation in all measures” di- 
rected towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepidoptera. All mem- 
bers receive the Journal and the News of The Lepidopterists’ Society. Prospective members should 
send to the Assistant Treasurer full dues for the current year, together with their full name, address, 
and special lepidopterological interests. In alternate years a list of members of the Society is issued, 
with addresses and special interests. 


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Send remittances, payable to The Lepidopterists’ Society, to: Ron Leuschner, Asst. Treasurer, 1900 
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Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepi- 
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Cover illustration: the Large Yellow Underwing, Noctua pronuba L., a newcomer to Connecticut 
during 1993, and a “missing person” in the recent Peterson’s Field Guide to Eastern Moths. This 
palearctic noctuid has rapidly expanded its range and population size throughout eastern North 
America since its introduction into Canada nearly two decades ago. Original pen and ink drawing by 
John Himmelman, 67 Schnoor Road, Killingworth, Connecticut, 06419, USA. 


VOme NAL. OF 
Hou LEPIDOPTERISTS’ SOCIETY 


Volume 51 1997 Number 3 


Journal of the Lepidopterists’ Society 
51(3), 1997, 197-207 


MALE MATE-LOCATING BEHAVIOR AND YEARLY 
POPULATION CYCLES IN THE SNOUT BUTTEREBLY, 
LIBYTHEANA BACHMANII (LIBYTHEIDAE) 


RONALD L. RUTOWSKI, BARBARA TERKANIAN, OFER EITAN 


Department of Biology, Arizona State University, 
Tempe, Arizona 85287, USA 


AND 


ANDREA KNEBEL 


Universitat Bielefeld, Verhaltensforschung, 
Postfach 100131, 33501 Bielefeld, Germany 


ABSTRACT. This paper describes mating behavior and seasonal changes in popula- 
tion size in the snout butterfly, Libytheana bachmanii. At a central Arizona study site, we 
found a dramatic peak in the abundance of snout butterflies in late May and early June, 
with a smaller peak in the fall. Both peaks lasted several weeks and were separated by pe- 
riods when few or no adult butterflies were found. Males are classic patrollers and search 
for females in and around the larval foodplant, desert hackberry (Celtis pallida). Courtship 
is like that of many other butterflies, with no distinctive displays by the male or female. 
We compare these results to those for the desert hackberry butterfly, Asterocampa leilia, 
which uses the same larval foodplant but has very different mate-locating tactics and, as 
some hypotheses predict, relatively stable and medium density populations from season to 
season. 


Additional key words: desert hackberry, central Arizona, courtship. 


The behavior of male insects at mate encounter sites varies along sev- 
eral axes. These include the time of day males visit sites, the duration of 
visits to a site, and whether males defend sites (Thornhill & Alcock 
1983). Ultimate explanations for differences within and among species 
in these components of male behavior at encounter sites invoke ecolog- 
ical variables such as population density and encounter site size and dis- 
tribution (Thornhill & Alcock 1983, Bradbury 1985). A special opportu- 
nity to study the ecological correlates and causes of mating systems 
arises when two or more species of similar size use the same encounter 


198 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


sites but differ in mate-locating tactics. In these situations, certain fea- 
tures of habitat and scale are naturally controlled, which permits a focus 
on differences in other factors such as population density that may have 
influenced the evolution of male mate-locating behavior. 

In the upper Sonoran Desert of western North America, two butterfly 
species similar in size occupy the same locale and use the same larval 
food plant, desert hackberry (Celtis pallida Torrey; Ulmaceae). These 
butterflies appear to exhibit striking interspecific differences in male 
mate-locating behavior and in population dynamics. In one of the spe- 
cies, the desert hackberry butterfly, Asterocampa leilia (Edwards) 
(Nymphalidae), males are classic “perchers” (Scott 1974) that occupy 
and defend perch sites on or next to the larval food plant (Austin 1977, 
Rutowski & Gilchrist 1988, Rutowski et al. 1991). A male may occupy 
the same site for several mornings. In contrast, males of the other spe- 
cies, the snout butterfly, Libytheana bachmanii Strecker (Libytheidae) 
have been described as perchers by some (Scott 1986) and “patrollers,” 
that conduct aerial searches within hackberry trees but do not occupy or 
defend perches, by others (Rutowski 1991). 

Selection should favor males that patrol when the costs of site tenac- 
ity and defense outweigh the benefits (Brown & Orians 1970, Rutowski 
1991). This should occur when population densities are extremely high 
or low. At high population densities the rate of interactions with intrud- 
ers should place a high cost on site defense; at low population densities, 
the low rate of female arrival will yield low returns from site defense. In- 
traspecific switches from perching to patrolling have been related to in- 
creases (Alcock & O'Neill 1986, Wickman 1988) and decreases (Wick- 
man & Wiklund 1983) in population density in other butterflies. The 
snout butterfly is notorious for undergoing large population explosions 
(“usually in late summer” Bailowitz & Brock 1991; “emigratory flights of 
millions of butterflies” Pyle 1981; Scott 1986), but the timing and dura- 
tion of these events and population sizes between outbreaks have not 
been quantitatively documented. Populations of A. leilia are apparently 
subject to much less fluctuation (Rutowski et al., 1996). 

To determine if an association exists between population characteris- 
tics and male behavior, we quantified population dynamics and features 
of the male and female behavior in L. bachmanii. Here we address the 
following questions: (1) what annual changes occur in the population 
size of this species; (2) what search tactics do males employ, and how do 
they differ from those of A. leilia; (3) what is the nature of courtship in 
this species; and (4) what is the mating history of individual females? Fi- 
nally, we compare our profile of L. bachmanii with previous data on A. 
leilia and relate this information to hypotheses about the role of popula- 
tion density in the evolution of male behavior at mate encounter sites. 


VOLUME 51, NUMBER 3 199 


METHODS 


Study site. We studied L. bachmanii at Round Valley in the Syca- 
more Creek flood plain, approximately 70 km NW of Phoenix, Arizona. 
The primary vegetation at this site includes shrubs and low trees, such 
as desert hackberry, mesquite (Prosopis spp.), catclaw (Acacia greggii 
Gray; Leguminosae), and paloverde (Cercidium spp.). The observations 
reported here were made from 1987 to 1995. 

Census techniques. To assess yearly and daily changes in popula- 
tion size of L. bachmanii, we selected and mapped a 270 m census trail 
in the study site. The trail ran through a stand of hackberry trees and 
was used previously to census A. leilia populations (Rutowski et al., 
1996). On each census, an observer walked the entire length of the route 
and recorded the location and behavior (perched or flying) of each L. 
bachmanii individual (males and females) seen on the census route. We 
tried to complete each census within 10 to 15 min to avoid counting mov- 
ing individuals more than once; the cost of this approach was that we 
could not reliably distinguish males and females on the wing, so they were 
not scored separately. We censused the population in this way on sunny 
days every seven to 14 days during the flight season and once a month at 
other times, for a total of 40 days, from March 1993 to October 1994. On 
35 of these days we ran four hourly censuses, from 0900-1200 h (MST). 
The other five days were during periods when no butterflies were found, 
so we ran only two or three censuses. At the end of each census, we mea- 
sured the air temperature in the shade at 1 m above the ground. 

Behavioral observations. Intrasexual and intersexual interactions 
were described from field observations. All interactions began when a 
male approached a conspecific. We recorded the durations of interac- 
tions from when an approaching male arrived within a few cm of the 
other individual until the interacting pair separated for the last time. 
Also, for a sample of perched males, we measured the height of each oc- 
cupied perch. Observations and video records of field interactions be- 
tween males and virgin females (reared from larvae collected in the 
field) of L. bachmanii were used to develop the description of courtship. 
To estimate mating frequencies of females, we dissected the females that 
had been collected at random and freeze killed during May, June, and 
July 1994. The abdomen of each was opened under insect Ringer's solu- 
tion, and the number of spermatophores in the bursa copulatrix counted. 

Tenure in hackberry trees. We attempted to determine if males 
behave differently in hackberry trees than they do in other trees. We se- 
lected a hackberry tree and a mesquite tree of similar size (about 3 m 
high and wide) that were representative of trees within the study site. 
An observer was stationed at each tree. At the same time, each observer 


200 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


—&— L. bachmanii 
—O— Temperature 


Max. number of L. bachmanii observed 
(Do) aunyesodwis | 


Mar. 93 Jun. 93 Oct. 93 Jan. 94 Apr. 94 Jul. 94 Oct. 94 
Date 


Fic. 1. Variation in the size of the L. bachmanii population and in the air temperature 
at 1 m with time of year. For each day on which we censused the population, the graph 
shows the maximum number of individuals observed on a single census and the tempera- 
ture at the time of that census. L. bachmanii abundance is dependent on season but not 
on temperature (see text for details). This figure includes data from five days in which only 
two or three censuses were run and no butterflies were seen. 


recorded the total time spent within the tree’s perimeter for each male 
L. bachmanii that arrived at the tree. The data were collected on two 
sunny days in May 1986, using two different pairs of trees. 


RESULTS 


Population dynamics. The number of L. bachmanii seen along the 
census route on any given day fluctuated dramatically within each year 
(Fig. 1). On most days the maximum number of butterflies seen on any 
given census did not exceed 10 individuals; however, on three out of 40 
census days we counted more than 60 individuals in at least one census. 
Flight seasons occurred at the study site twice each year: over six to 
eight weeks during May and June, and from four to six weeks during 
September and October. Peak population density was much greater dur- 
ing the spring flight season than in the fall. The average number of indi- 
viduals observed on a day varied significantly with time of year (Kruskal 
Wallis test, n = 40 days, P = 0.006; Fig. 1). However, the average num- 
ber seen on a day when the butterflies were active did not covary with 
air temperature at | m at noon on that day (Kruskal Wallis test, n = 35 


VOLUME 51, NUMBER 3 201 


% flying 


0 20 40 60 80 100 
L. bachmanii observed per census 


Fic. 2. The proportion of individuals seen in flight (rather than perched) as a function 
of population density (n = 1222). Butterflies that changed from flying to perching (or vice 
versa) while being censused not included. 


days, P = 0.437). Although L. bachmanii are most abundant when ambi- 
ent temperature is between 30 and 40°C, there were days on which 
these temperatures occurred but no butterflies were seen (Fig. 1). 

Whether L. bachmanii were seen perched or flying during a census 
was related to the density of butterflies (Fig. 2). When we counted more 
than 20 butterflies along the census trail, at least 80% of them were fly- 
ing. However, when counts were 20 or lower, the proportion of individu- 
als in flight was variable, ranging from 0 to 100% of butterflies observed. 

No consistent pattern of daily activity was found in the mornings (be- 
tween 0900-1200 h). Time of day did not explain at all the number of 
adults seen on a census on days when the butterflies were active 
(ANOVA with time nested in date on log transformed counts during 
flight seasons, df = 75, P = 0.999). Moreover, we saw no consistent 
movement patterns that would lead us to describe these populations as 
migratory, as previous accounts have done (e.g., Pyle 1981). 

Along the census route, snout butterflies were most commonly found 
near hackberry trees, their larval foodplant. When we divided the cen- 
sus route into 5 m segments and totaled butterfly sightings for each seg- 
ment, we found that butterflies were not uniformly distributed along the 
trail (Fig. 3; c? = 1034, df = 53, P < 0.0001). Similarly, we found a non- 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


140 
120 
100 
80 
60 


40 


Total number of L. bachmanii observed 


20 


0 25 50 75 100 125 150 17h5) 200 225 250 


Location along the census route (m) 


Fic. 3. Spatial distribution of L. bachmanii sightings (n = 1797) along the census 
route. The number observed on each 5 m segment is the total observed over 140 censuses 
on 35 days. 


uniform distribution of hackberry trees along the trail (c2 = 36.9, df = 
53, P < 0.05). However, there was a significant, positive correlation be- 
tween the number of L. bachmanii observed in each 5 m section of the 
census route and the number of hackberry trees in a section (Fig. 4; r = 
0.56, df = 53, P < 0.0001). 

Male searching behavior. Males were seen flying and perched in 
the study area, and flying males moved both within and among hack- 
berry trees. Within hackberry trees, males moved slowly up and down 
close to branches and leaves. Perched butterflies were approached and 
inspected. Both flying and perching males approached and chased other 
butterflies (both conspecific and heterospecific) that flew nearby; how- 
ever, at times two or three butterflies perched within a few centimeters 
of each other. Interactions with conspecific males in the spring flight pe- 
riod lasted an average of 14.6 + 13.5 sec (median = 9.7 sec, range = 
3.9-43 sec, n = 8). Males inspected heterospecifics for an average of 
12.7 = 110.3 see (median = ‘7/3 sec, range — 2.8-36-2 sec, 1916) aiaie 
interaction durations were not significantly different (Mann-Whitney U 
test, P > 0.9). 

Flying males spent significantly more time in hackberry trees than 
they did in mesquite (Fig. 5; Wilcoxon rank sum test, n = 22, P = 0.001). 


VOLUME 51, NUMBER 3 203 


140 

Oo 

S 120 

cb} 

” 

zs) 

= 100 

Cc 

140} 

5 

So 80 

© 

Q 

a 

“5 60 

= 

cb) 

2 

: 40 

(= 

To 

ie) 

pati 20 
0 

0 1 2 3 4 


Hackberry trees in 5m segments 


Fic. 4. The relationship between the number of hackberry trees and the number of L. 
bachmanii sightings in 5 m segments along the census route. There is a significant positive 
correlation between these variables (n = 1797, r = 0.56, df = 53, P < 0.0001). 


Males moved through hackberry trees in an erratic, zigzag fashion. In 
contrast, their flight through mesquite trees was more direct, causing 
them to pass through the tree more quickly. 

Males perched on exposed twigs and branches. Average perch height 
was 1.92 + 0.31 m (median = 1.87 m, range = 1.5—2.5 m, n = 17) above 
the ground. The body of a perched male was oriented upwards with the 
wings closed or partially opened. The average time a male spent at a 
perch on a typical summer day was 77 + 90.6 sec (range 3.4—430.3 sec, 
n = 55). 

Courtship and mating behavior. Courtship leading to copulation 
began when a male approached a perched or flying female. A flying fe- 
male was chased by a male until she alit on vegetation. A perched re- 
ceptive female usually remained still with her wings folded as a male ap- 
proached. The male then alit behind the female and moved alongside her 
with his head oriented in the same direction as hers. He then curled his 
abdomen toward the female, probing between her hindwings until he at- 
tained genital contact. After coupling, the male tumed to face away from 
the female. During copulation, some females took flight, carrying the 
male with his wings closed, suspended head-down from her abdomen. 


204 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


] Hackberry 
Ha Mesquite 


Males observed 


0 2 4 6 8 10 12 14 16 18 20 >20 


Time in tree (Sec.) 


Fic. 5. Time spent by males in hackberry trees versus mesquite trees. See text for 
details. 


Copulation ended when the pair uncoupled. Four pairs were found in 
copula, and the average time until they separated was 88.5 + 40 min 
(range = 45-140 min, n = 4). During copulation the male passed a sper- 
matophore to the female. The females collected for spermatophore 
counts were fresh to slightly worn in condition and carried an average of 
1.28 + 0.66 spermatophores (n = 28, range = 1—4 spermatophores). 
When females carried more than one spermatophore, one was roughly 
teardrop shaped, approximately twice as long as wide, and all others 
were partially or completely flattened and depleted. This suggests that 
matings do not occur in rapid succession. After copulation females flew 
about searching for oviposition sites. Eggs were laid singly, often on the 
end of sprigs of new hackberry growth. 

Unsuccessful courtships lasted an average of 44.6 + 79.7 sec (median 
= 42.3 sec, range = 23-78 sec, n = 6). In these interactions, flying fe- 
males either did not alight, alit and fluttered their wings, or suddenly 
dropped toward the ground and alit. In the latter case, the male might 
search for the female, but he left after a brief time if he did not relocate 
her. Perched females may hinder males by spreading their wings and el- 
evating their abdomens or by taking flight. On one occasion a male pur- 


VOLUME 51, NUMBER 3 205 


sued a female in direct upward flight for about 5 m. The chase lasted for 
less than one minute, until one of the butterflies was captured by a cliff 
swallow (Hirundo pyrrhonota Vieillot). 


DISCUSSION 


The major results of this study are as follows. First, the L. bachmanii 
population that we studied in central Arizona displays an explosive in- 
crease in numbers twice a year, especially in the late spring. The reasons 
for these dramatic changes are not known. Apparently the snout butter- 
flies diapause once or twice a year, but the stage in the life history at 
which this occurs or why it occurs at these times are not known. One 
possible answer is that population cycles are tied to the phenology of the 
desert hackberry. Hackberry trees produce new vegetation after the 
rainy periods that occur regularly each winter and mid to late summer. 
The new vegetation may foster the development of the larvae. 

Second, both males and females are found primarily in or near the lar- 
val foodplant. Females are seeking oviposition sites; males are seeking fe- 
males. Our observations show that males entering hackberry trees 
change their behavior from direct flight to a zigzag searching of the veg- 
etation. We conclude that males are seeking both newly emerged virgin 
females and previously mated females. Newly-emerged females are likely 
to be common in hackberry trees because the larvae pupate on the larval 
foodplant (pers. obs.). In addition, we have in several instances seen a 
male mated with a newly emerged female (wings still flexible) next to a 
pupal exuvium on hackberry. Males are also probably looking for previ- 
ously mated females who are ovipositing but are ready to mate again. 
Spermatophore counts show that females will mate more than once. Use 
of the larval foodplant as the mate encounter site has been reported for 
other butterflies (Lederhouse et al. 1992, Rutowski & Gilchrist 1988). 

Third, at the larval foodplant, males use primarily a patrolling strategy 
to locate females. Figure 2 suggests the incidence of a territorial strat- 
egy at low population densities; however, several observations argue 
against this. Males always show low interest in other males and little site 
tenacity. In addition, we never saw any male-male contacts that led to 
the spiraling aggressive interactions described for some territorial spe- 
cies (e.g., Baker 1972). Finally, males and females are not distinguished 
in Fig. 2, and temperature, which could also play a role, is not con- 
trolled. Even if we ignore these confounding variables, most individuals 
at all population densities are on the wing. 

Fourth, although the snout butterflies are a worldwide family of only 
nine species reported to have diverged from other butterfly lineages at 
least 35 million years ago (Emmel et al. 1992) and have unique palpal 
and wing morphologies, their courtship and mating behavior are very 


206 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


much typical of that reported generally for butterflies (Scott 1973, Sil- 
berglied 1978, Drummond 1984, Rutowski 1984). 

Relationship between population density and male behavior at 
encounter sites. The documentation of the dramatic seasonal changes 
in population density in L. bachmanii provided here supports the hy- 
pothesis that population density is an important determinant of male 
tactics. Because population densities are usually extremely low or, for a 
brief period, extremely high, the costs of site tenacity and defense 
should outweigh the benefits. Patrolling has then evolved as the primary 
mate-locating tactic in this species. 

The idea that population density is an important determinant of male 
behavior at encounter sites is further supported by comparing the re- 
sults reported here for L. bachmanii with what is known of Asterocampa 
leilia, the desert hackberry butterfly. Males in this species also use 
desert hackberry as the mate encounter site. However, they occupy and 
vigorously defend perching sites on or next to hackberry trees where 
they sit and wait for females to fly by. In contrast to L. bachmanii and in 
support of the population density hypothesis, the population of A. leilia 
at the Round Valley site is relatively stable for a long period each year. 
On the same census route used in this study, we found an average of 
about 10 perching males from April to November (Rutowski et al., 
1996). Compared with L. bachmanii their populations are at an inter- 
mediate level and relatively stable, which has, in our view, favored the 
evolution site tenacity and defense. 3 

Meteorological variables have also been proposed to explain intra- 
and interspecific differences in whether males patrol or perch at en- 
counter sites (Dennis 1982, Wickman 1985, 1988, Alcock 1994). How- 
ever, in neither L. bachmanii (this study) nor A. leilia (Rutowski et al. 
1994) have we seen any evidence that males switch from patrolling to 
perching or vice versa with changes in season, temperature, or time of 
day. Also, both species engage in their respective mate locating activities 
at the same time of the day. In summary, while this study implicates dif- 
ferences in population density to explain interspecific differences in 
male behavior, other nonmutually exclusive explanations such as tem- 
perature and predation will also need to be examined. 


ACKNOWLEDGMENTS 
We thank K. Wallace, K. Fales, N. Compton, and S. Pitnick for stimulating discussions 
and help in the field. This work was supported with funds from NSF Grant No. 83-00317 
to RLR and from the College of Liberal Arts and Sciences at Arizona State University. 


LITERATURE CITED 


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Nymphalidae). Ethology 97:103—-118. 

ALCOCK, J. & K. M. O'NEILL. 1986. Density-dependent mating tactics in the grey hair- 
streak, Strymon melinus (Lepidoptera: Lycaenidae). J. Zool. Lond. A 209:105—113. 


VOLUME 51, NUMBER 3 207 


AUSTIN, G. T. 1977. Notes on the behavior of Asterocampa leilia (Nymphalidae) in south- 
ern Arizona. J. Lepid. Soc. 31:111-118. 

BAILOWITZ, R. A. & J. P. BRocK. 1991. Butterflies of southeastern Arizona. Sonoran 
Arthropod Studies, Inc., Tucson, Arizona. 

BAKER, R. R. 1972. Territorial behaviour of the nymphalid butterflies, Aglais urticae (L.) 
and Inachis io (L.). J. Anim. Ecol. 41:453—469. 

BRADBURY, J. W. 1985. Contrasts between insects and vertebrates in the evolution of male 
display, female choice, and lek mating, pp. 273—289. In Hélldobler, B. & M. Lindauer 
(eds.), Experimental behavioral ecology and sociobiology. Fisher, New York. 

BRowN, J. L. & G. H. OrRIANS. 1970. Spacing patterns in mobile animals. Ann. Rev. Ecol. 
Syst. 1:239—262. 

DENNIS, R. L. H. 1982. Mating location strategies in the wall brown butterfly, Lasiommata 
megera (L.) (Lep., Satyridae): wait or seek? Entomol. Record 94:209—214; 95:7-10. 

DRUMMOND, B. A., III. 1984. Multiple mating and sperm competition in the Lepidoptera, 
pp. 291-370. In Smith, R. L. (ed.), Sperm competition and the evolution of animal 
mating systems. Academic Press, New York. 

EMMEL, T. C., M. C. MINNO & B. A. DRUMMOND. 1992. Florissant butterflies: a guide to 
the fossil and present-day species of central Colorado. Stanford University Press, 
Stanford, California. 148 pp. 

LEDERHOUSE, R. C., S. CODELLA, D. W. GROSSMUELER & A. D. MACCARONE. 1992. Host 
plant-based territoriality in the white peacock butterfly, Anartia jatrophae (Lepi- 
doptera: Nymphalidae). J. Insect Behav. 5:721—727. 

PYLE, R. M. 1981. The Audubon field guide to North American butterflies. Alfred A. 
Knopf, New York. 916 pp. 

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Res. Lepid. 23:125—142. 

. 1991. The evolution of male mate-locating behavior in butterflies. Amer. Nat. 
LS 11211139) 

Rutowski, R. L., M. J. DEMLONG & B. TERKANIAN. 1996. Seasonal variation in male 
mate-locating behavior in the desert hackberry butterfly, Asterocampa leilia. J. Insect 
Behav. 9:921—931. 

RuTowskI, R. L., M. J. DEMLONG & T. LEFFINGWELL. 1994. Behavioural thermoregula- 
tion at mate encounter sites by males butterflies (Asterocampa leilia, Nymphalidae). 
Anim. Behav. 48:833—841. 

RuTOWSKI, R. L., J. L. DICKINSON & B. TERKANIAN. 1991. Behavior of male desert hack- 
berry butterflies, Asterocampa leilia (Nymphalidae) at perching sites used in mate lo- 
cation. J. Res. Lepid. 30:129-139. 

RuTowskKI, R. L. & G. W. GILCHRIST. 1988. Male mate-locating behavior in the desert 
hackberry butterfly, Asterocampa leilia (Nymphalidae). J. Res. Lepid. 26:1—12. 

ScorTT, J. A. 1973. Mating of butterflies. J. Res. Lepid. 11:99—127. 

. 1974. Mate-locating behavior of butterflies. Amer. Midl. Nat. 91:103—117. 

. 1986. The butterflies of North America. Stanford University Press, Stanford, Cal- 
ifornia. 583 pp. 

SILBERGLIED, R. E. 1977. Communication in the Lepidoptera, pp. 362—402. In Sebeok, 
T. (ed.), How animals communicate. Indiana University Press, Bloomington. 

THORNHILL, R. & J. ALCOCK. 1983. The evolution of insect mating systems. Harvard Uni- 
versity Press, Cambridge. 547 pp. 

WICKMAN, P.-O. 1985. The influence of temperature on the territorial and mate locating 
behaviour of the small heath butterfly, Coenonympha pamphilus (L.) (Lepidoptera: 
Satyridae). Behav. Ecol. Sociobiol. 16:233-238. 

. 1988. Dynamics of mate-searching behaviour in a hilltopping butterfly, Lasiom- 
mata megera (L.): the effects of weather and male density. Zool. J. Linn. Soc. 
Sts ras ie 

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speckled wood butterfly (Pararge aegeria). Anim. Behav. 31:1206—1216. 


Received for publication 24 March 1996; revised and accepted 3 June 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 208-217 


RELATEDNESS AND GREGARIOUSNESS IN THE ORANGE- 
STRIPED OAKWORM, ANISOTA SENATORIA (SATURNIIDAE) 


ADAM H. PORTER!, SEAN J. CADARET, SCOTT A. JOHNSON, 
HIDETAKA MIZOHATA, ANDREA I. BENEDETTER?, 
CATHLEEN L. BESTER, JENNIFER L. BORASH, SCOTT D. KELLY, 
GRETCHEN S. BUEHNER, AND MARILYN L. SHERMAN 


Department of Biological Sciences, Bowling Green State University, 
Bowling Green, Ohio 43403, USA 


ABSTRACT. Oakworm larvae live in groups and provide a convenient model system 
for the study of gregarious behaviors of caterpillars. Most caterpillar groups consist largely 
of related individuals, so the costs and benefits of gregariousness must be considered 
within the framework of kin selection. In this study, we use allozymes to estimate related- 
ness within 11 groups of 4th—5th instar larvae. Allozyme diversity was high in four marker 
loci, with an average heterozygosity of H_,,,, = 0.376 + 0.029 (SD) when frequencies were 
pooled over groups. We found a relateaniece of r = 0.31 + 0.056, a relatively low value 
given that full-siblings show r = 0.5. The mean nearest-neighbor distance among plants 
was 4.34 + 3.48 m, and among groups it was 5.25 + 4.15 m. Within the framework of a 
mathematical dispersal model calibrated using observations of larval movement in the 
field, these values suggest that only about 4% of wandering larvae ever find new hosts, and 
that exchange is negligible among groups on different plants. Adult behaviors, either mul- 
tiple mating or aggregation of egg clutches, are therefore probably responsible for the ob- 
served relatedness values. 


Additional key words: caterpillar ecology, Lepidoptera, dispersal, group living. 


Oakworm moths (Anisota senatoria J. E. Smith) are primarily known 
by the considerable destruction their gregarious larvae can cause to the 
foliage of oaks in eastern North America (e.g., Coffelt, Schultz & Wolf 
1993). The larvae are brightly colored and easily manipulated in the 
field, making them excellent models for the study of communal behav- 
iors of caterpillars. 

Natural history data relevant to the communal behavior of A. senato- 
ria larvae are available from recent studies to assess and control their 
impact as pests in suburban landscapes (Coffelt & Schultz 1990, 1993). 
Adults emerge in late June to early July and copulate diurnally (Fergu- 
son 1971), and females lay clutches of 200-700 eggs on the undersides 
of oak leaves (Coffelt & Schultz 1990). Host plants in other families 
have been reported in the older literature (compiled in Teitz 1972), but 
these are probably erroneous (Ferguson 1971). Females are weak fliers 
and over 90% of the clutches are laid within 5 m of the ground (Coffelt 


‘Corresponding author; current address: Dept. Entomology, University of Massachu- 
setts, Amherst, Massachusetts 01003, USA, email:aporter@ent.umass.edu 
2Institut fiir Zoologie, Universitat Salzburg, Hellbrunnerstr. 34, A-5020 Salzburg, Austria 


VOLUME 51, NUMBER 3 209 


& Schultz 1994). The larvae appear to be confined to older leaves (Law- 
son et al. 1982, Coffelt, Schultz & Banko 1993). The larvae feed gregar- 
iously, resting on stems and twigs and relocating to nearby (often higher) 
branches as defoliation proceeds (Coffelt & Schultz 1994). Mature lar- 
vae pupate in the soil beneath their host trees. Most populations are 
univoltine, but bivoltine populations occur in the southern portion of 
the range (Ferguson 1971, Coffelt & Schultz 1994). 

In most non-eusocial insects studied, group-living is initiated when fe- 
males lay clutches of eggs (e.g., the eastern tent caterpillar, Malacosoma 
americanum (F.) (Lasiocampidae) (Costa & Ross 1993); the willow leaf 
beetle, Plagiodera versicolora Laicharting (Chrysomelidae) (McCauley 
et al. 1988)). Group living may subsequently be promoted by larval be- 
haviors that maintain and enhance gregariousness, as in M. americanum 
(Fitzgerald & Peterson 1988) and Pryeria sinica Moore (Zygaenidae) 
(Tsubaki 1981), or it may be maintained simply by the absence of larval 
dispersal from the oviposition site. 

Several selection pressures may act simultaneously on group-living 
behaviors of larval insects. Factors acting against group living in cater- 
pillars include greater risks of detection by predators (Morris 1972, 
Stamp & Bowers 1988) and parasitoids, increased cannibalism, and 
greater chances of intragroup competition via defoliation of the host 
plant (Tsubaki & Shiotsu 1982). Those favoring gregariousness include 
facilitation of feeding on tough host leaves (Ito et al. 1982; and sug- 
gested for early instars of A. senatoria by Hitchcock 1961), the sharing 
of silken trails and nests (Fitzgerald & Peterson 1988), enhanced ther- 
moregulation at low air temperatures (Stamp & Bowers 1990a), and po- 
tentially greater effectiveness of defensive behaviors against parasitoid 
attack (Stamp & Bowers 1990b). Additional benefits of larval gregari- 
ousness include enhanced effects of chemical defenses (Tostowaryk 
1972 for Neodiprion sawfly larvae) and warning coloration (Sillen-Tull- 
berg 1990) for distasteful species, and selfish-herd effects mediated by 
predation, provided that predation is not too intense (Hamilton 1971). 
These selection pressures may change with age, influencing the degree 
of grouping and cooperation that is optimal for larvae of different ages 
(Cornell et al. 1987, McCauley et al. 1988, Costa & Ross 1993). Inas- 
much as individuals in larval groups are likely to be related because of 
maternal oviposition behavior, the costs and benefits of gregariousness 
need to be interpreted within a kin-selection framework. 

In this study, we use allozymes to assess relatedness within late-instar 
groups of A. senatoria larvae, and present results of a simple experiment 
on larval dispersal ability to assess its influence on relatedness. Other 
factors that may influence A. senatoria’s gregariousness, outlined above, 
will be pursued in separate studies. 


210 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


MATERIALS AND METHODS 


Fourth and fifth instar larvae of A. senatoria were observed on indi- 
vidual red oaks (Quercus rubra L.; Fagaceae) at the Girdham Road sand 
dunes in Oak Openings Metropark in Lucas Co., Ohio, on 9 September 
1995. The site includes a mixture of mature, sapling, and seedling age 
classes on a sandy substrate, and larval groups were abundant and easily 
accessible on the smaller plants. Larvae were identified using the key in 
Ferguson (1971). Few groups were seen at heights >5 m, in agreement 
with Coffelt & Schultz (1994). 

Relatedness. Eleven larval groups were arbitrarily chosen and 
from each, 15 individuals (or all individuals if <15 were present) were 
removed and frozen at —80°C. Tissue was sampled by shaving slices 
from the abdomens of the still-frozen larvae, taking care not to include 
gut contents. Starch gels were prepared using standard techniques 
(Porter & Matoon 1989). Seven loci were stained: glutamic-oxaloacetic 
transaminase (GOT-1, GOT-2), malate dehydrogenase (MDH-1, MDH- 
2), malic enzyme (ME), phosphoglucomutase (PGM), and phosphoglu- 
cose isomerase (PGI). Alleles were scored alphabetically by locus. Indi- 
viduals with rare alleles were re-run in adjacent lanes to confirm their 
scoring, and unresolved individuals were re-run as well. 

The relatedness statistic, r, describes the extent to which individuals 
within groups share alleles, beyond the degree to which alleles are 
shared with an ‘average’ individual in the population as a whole. Grafen 
(1985) provides an excellent discussion of the interpretation of related- 
ness statistics. Relatedness was calculated using equation 6 of Queller & 
Goodnight (1989), a method that accounts for sampling bias. All calcu- 
lations were performed using a population genetic analysis program 
written by A. H. Porter, which is available upon request. 

Dispersal. Larvae were tested to determine their dispersal and re- 
aggregation capabilities following disturbances. Four plants <2 m in 
height were found with larval groups (n = 22, 4, 5, 6) on them. Concen- 
tric circles with 1, 2, and 3 m radii were drawn in the sand around these 
plants and the branch containing the group was jolted to simulate the 
arrival of a potential vertebrate predator. The larvae dropped to the 
ground and their movement distances and directions were monitored at 
2 min intervals for 10 min. These were converted to average movement 
rates to provide a rough estimate of the ability of larvae to move among 
plants. The circle was divided into four quadrats and movement direc- 
tion was determined for each larva as the quadrat it occupied after 10 
min. Larvae could disperse in any direction so circular statistics 
(Batschelet 1981) were used on these data to assess whether the larvae 
tended to move together. 


VOLUME 51, NUMBER 3 


TABLE lL. 


ber of stainable individuals for each locus. 


allele 


GOT-1 


MDH-2 


PGI 


211 


Allele frequencies (s.e.) of the pooled data. Sample sizes (n) indicate num- 


PGM 


n 107 153 158 142 

A 0.009 (0.001) 0.761 (0.002) 0.066 (0.001) 0.025 (0.001) 
B 0.944 (0.001) 0.239 (0.002) 0.025 (0.001) 0.056 (0.001) 
C 0.047 (0.001) 0.051 (0.001) 0.486 (0.003) 
D 0.725 (0.002) 0.419 (0.003) 
E 0.133 (0.001) 0.014 (0.001) 


Distances among hostplants and larval groups. To estimate the 
probabilities of larvae moving between plant or to new groups, host- 
plants and all larval groups within 2 m of the ground were first identi- 
fied and mapped within a 50 x 100 m area. This area was chosen because 
it contained sufficient plants and larval groups for a statistical analysis, 
and it was set apart from other areas with infested plants. The map was 
constructed by creating a lattice of triangles with sampled plants at the 
vertices, then measuring the distances between plants; nine missing 
measurements were estimated from field notes and sketches. Under this 
system, the coordinates of a third point of a triangle can be found using 
simple geometry once the first two are established. The lattice was 
thereby converted to a Euclidean coordinate system by establishing the 
coordinates of the first two points along a north-south line (the first 
point is at (0, 0), the second is at (0, d), where d is the distance between 
points) and iterating through the lattice until all coordinates were calcu- 
lated. The distances between any two lattice points can then be found 
directly using the Pythagorean relationship. All calculations were per- 
formed using a Mathematica notebook (v2.2; Wolfram 1991). This con- 
venient method provides an explicit map of all points of interest on the 
site without the need for erecting a grid. The method readily yields the 
distributions of interplant and intergroup distances, information that is 
not obtainable using nearest-neighbor measurements. 

We estimated the proportion of wandering larvae that find new plants 
from the geometry of the plants in the study area. The angle subtended 
by a plant at distance d is 0.2/d radians, and the proportion of a search 
circle that culminates in a plant is 0.1 a! XZ d-!. When plant density is 
low, these proportions are rather small, and to provide a conservative as- 
sessment of the role of dispersal, we chose assumptions that overesti- 
mated the chances of concluding that a larva would find a new plant. We 
thus assumed a relatively constant heading for a searching caterpillar, 
and that any plant within 10 cm of its path would be detected. A con- 
stant heading is the best strategy for a searching caterpillar when plant 
density is low (Jones 1977). As this assumption is relaxed and the 


212 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


to another hosiplant 


relative frequency 
S99 © 
Oo f O 


© 
LS) 


to another group 


= 
= 


0 1 2 3 4 3) 6 7 
traveling time (hrs) 


Fic. 1. Estimated distributions of traveling times for larvae successfully dispersing be- 
tween plants, and between groups on separate plants. An additional 96% of larvae would 
not find plants at all. 


propensity increases for larvae to change their heading during their 
search, they will wander along increasingly longer average paths before 
reaching new hosts. This is equivalent to increasing the distances be- 
tween plants under a constant-heading search, and results in a some- 
what lower probability of finding a new host. Dethier (1989) found that 
dispersing caterpillars tend to maintain relatively straight paths unless 
perturbed by obstacles. Our study plants were chosen for the relatively 
few obstacles in the sandy substrate around them, and this too may yield 
relatively generous estimates of dispersal capabilities. The value of 10 
cm for the radius of detection was also chosen to be generous. When 
unaided by silken trails of conspecifics, other caterpillars species tested 
rarely showed orientation to hosts beyond 5 cm distance (Dethier 1959, 
Saxena & Khattar 1977, Saxena et al. 1977), though vertical stems (host 
or not) induced orientation in some species at 50 cm, even as far as 3 m 
(Doane & Leonard 1975, Roden et al. 1992). We did not adopt these 
higher values for three reasons. First, few larvae in our dispersal experi- 
ments returned to the source plant despite its close proximity, suggest- 
ing that such orientation capabilities are negligible in Anisota senatoria. 


VOLUME 51, NUMBER 3 NS} 


Second, plants of several non-host species were present in the study 
area in addition to young oaks, and orientation to these would tend to 
confound the search for appropriate hosts. Third, these higher values 
are based on orientation on a smooth substrate, and Dethier (1989) 
showed that as the substrate becomes more complex, search paths be- 
come more convoluted and larvae are increasingly apt to be influenced 
by encounters with obstacles at close range. 


RESULTS 


Genetic diversity and relatedness. The ME locus showed clear 
polymorphism, but because several alleles were difficult to resolve in 
heterozygotes, this locus was dropped from the analyses. MDH-1, the 
anodal locus, showed patterns that appeared to indicate polymorphism, 
but these were not repeatable and this locus was also omitted. GOT-2, 
the cathodal locus, showed no polymorphism. The remaining loci, GOT- 
1, MDH-2, PGI and PGM, showed sufficient polymorphism for use in 
the relatedness analysis. Allele frequencies of the pooled groups are 
shown in Table 1. The observed heterozygosity (s.e.) of the pooled groups 
was H,,,. = 0.386 (0.022), with expected heterozygosity of H,,,, = 0.376 
(0.029). All groups showed polymorphism at >1 locus. Taken together, 
these results indicate that there was sufficient polymorphism available 
for a robust relatedness analysis. 

The relatedness (SD) among late-instar A. senatoria group-mates was 
r = 0.31 (0.056), where the standard deviation is assessed by jackknifing 
over loci. This standard deviation drops to 0.033 if the variation is as- 
sessed over groups. 

Larval dispersal capabilities. Oak plants at our site were sepa- 
rated by a mean (s.d.) nearest-neighbor distance of 4.34 (3.48) m. We 
found nearest-neighbor larval groups in our site to be separated by 5.25 
(4.15) m (n = 18 groups), with the closest groups being on plants less 
than | m apart. 

Fallen larvae rarely went back to the same plant. Upon falling, they 
moved at variable rates at relatively constant headings, averaging 1.5 
(2.2) cm/sec. 32% of the larvae moved only little and thus did not leave 
the inner circle. There was no evidence that caterpillars took similar 
headings (Rayleigh tests, P > 0.05), indicating that they do not travel in 
groups once disturbed in this manner. 

Based on our dispersal model, only 4% of the larvae that began 
searching (i.e., the 68% of larvae that traveled at least 1 m in the first 10 
min) would ever encounter another host plant. This does not account 
for movement to plants off the grid, but these were mostly far enough 
away that the chances of a dispersing larva encountering them could be 
considered negligible. As noted in the Methods, this estimate is based 


214 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


on assumptions that are already somewhat generous, and though it is 
low, it is still probably an overestimate. The low probability of finding a 
new host should therefore impose a strong limit on the exchange among 
larval groups on different plants. 

We estimated the expected distribution of traveling times between 
plants by dividing each movement rate estimate by each distance be- 
tween plants on the grid. Most of the fallen larvae that do find new 
plants (Fig. 1) would arrive there within the first hour (mean = 62 + 78 
min), and almost all successful larvae would find a new plant within 4 hr. 
Because of the spatial relationships among plants with and without lar- 
vae, approximately 65% of these new plants would already have larval 
groups, so the distribution of traveling times among groups (mean = 57 
+ 75 min) is similar to that among plants (Fig. 1). Traveling time to new 
plants thus does not appear to be limiting exchange among groups, at 
least for the 4th and 5th instar larvae we studied. Larvae may also dis- 
perse of their own accord (Coffelt & Schultz 1993), whereupon they 
could expect to find suitable plants, and possibly join other groups, with 
similar probabilities and within similar time frames. 


DISCUSSION 


The degree of relatedness is moderate, well below the r = 0.5 value 
expected if larvae were always full siblings, but above the r = 0.25 level 
expected from half-sibs. This relatedness value is high enough to have 
strong effects on the realized costs and benefits of the gregarious behav- 
iors of the larvae (Hamilton 1964). There are several ways that this level 
of relatedness could be realized, and these may be divided among be- 
haviors of the ovipositing females and the subsequent behaviors of their 
larvae. 

Larval behaviors that may reduce intragroup relatedness include ac- 
tive wandering to new groups during foraging bouts, as seen in Malaco- 
soma americanum (Costa & Ross 1993), and inadvertent dispersal to 
new groups after falling off the host plant, as seen in Hemileuca lucina 
(Stamp & Bowers 1987). At Oak Openings, smaller plants with larval 
groups were often badly defoliated, and some had been abandoned by 
larvae. In some cases, we found small groups on plants with no evi- 
dence of the usual molt skins left by earlier instars, and these groups 
were adjacent to abandoned, defoliated plants. Even though host plants 
were relatively close together by human standards (4.34 m nearest 
neighbor distance), the distances remain formidable for wandering 
caterpillars, and only about 4% were likely to encounter new hosts. 
Plants with larval groups were still further apart, at 5.25 m, and we ex- 
pect only a negligible proportion of larvae to be exchanged among them. 
This implies that the reduction in relatedness below r = 0.5 on the small, 


VOLUME 51, NUMBER 3 215 


isolated plants in our study is attributable more to the behaviors of adult 
parents, rather than to wandering by the larvae. However, intergroup 
exchange and coalescence in A. senatoria may be much more likely 
among clutches laid on the same plant, as seen in Malacosoma (Costa & 
Ross 1993). We would also expect it to be more pronounced at high 
density and later instars, when the defoliation rate is highest and larvae 
are forced to wander, and when larvae are big enough to travel at the 
rates we observed. 

Adult behaviors that would reduce relatedness, not addressed in this 
study, include multiple mating with sperm-mixing that would result in 
broods of mixed full- and half-sibs (Wade 1982, 1985, McCauley & 
O'Donnell 1984), and the aggregation of egg clutches onto the same 
plant by different females (Wade 1985). These behaviors would be less 
effective in reducing relatedness to the extent that related adults mated 
or aggregated their clutches (Wade 1985). Of course, parental behaviors 
would influence the degree of relatedness and gregariousness initially 
displayed by larvae, but should not create subsequent changes in gregar- 
iousness associated with larval age. 


ACKNOWLEDGMENTS 


We thank Joseph Croy of the Toledo Metroparks system for the research permit for the 
Oak Openings Metropark, and the Biological Sciences department for funding the study. 
This project was undertaken as part of a graduate course, Methods in Ecology, Behavior, 
and Evolution, offered at BGSU. We thank S. Krueger, P. Passalacqua, & R. Schneider for 
their assistance with data collection, and E. Jakob, S. Vessey, W. Watt, and two anonymous 
reviewers for valuable comments on the manuscript. 


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VOLUME 51, NUMBER 3 DIT 


TsuBAKI, Y. 1981. Some beneficial effects of aggregation in young larvae of Pryeria sinica 
Moore (Lepidoptera: Zygaenidae). Res. Pop. Ecol. 23:156—167. 

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iors in diploid and haplo-diploid organisms. J. Theor. Biol. 95:351—368. 

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Received for publication 7 March 1996; revised and accepted 13 July 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 218-226 


BIOLOGY OF THE BLACK-ANTENNA RACE OF PHYCIODES 
THAROS THAROS (NYMPHALIDAE) IN ONTARIO 


PAUL M. CATLING 
2326 Scrivens Drive, R. R. #3, Metcalfe, Ontario KOA 2P0, Canada 


ABSTRACT. The black-antenna race of P. tharos tharos (Drury) occurs north to the 
Canadian zone of the Ottawa valley, far beyond the Carolinian or Upper Austral range 
limit previously indicated for the species. It is also reported for the first time from north- 
ern New York State and Quebec. The northernmost colonies are restricted to alvars or old 
pastures over limestone, but hay field and old field mosaics, dune slacks and prairies on 
sandy soils are utilized farther south. The foodplant at the northernmost sites was Aster 
ciliolatus based oviposition in the field and subsequent rearing in the laboratory. Available 
evidence suggests that the black-antenna race of P. tharos tharos may have invaded the 
Ottawa valley over the past five years: it is now widespread but local in southern Ontario, 
having been found in 65 locations. The flight period extends from late May to late Octo- 
ber and is essentially continuous at some locations. 


Additional key words: oviposition, foodplants, Aster ciliolatus, habitat, flight period. 


Recent books (e.g., Scott 1986a, Opler & Krizek 1984, Opler & Ma- 
likul 1992) and monographs (Scott 1986b, 1994) follow Oliver (1980) in 
splitting P. tharos into two entities: the northern P. cocyta (Cramer) 
(= P. selenis, P. pascoensis, P. morpheus) and the more southern P. 
tharos tharos (Drury). Unfortunately, ecological and distributional data 
on the two species are confused because they previously were treated as 
one, and it is often unclear to which species certain information applies. 
The purpose of the present work is to provide ecological, distributional, 
and behavioral data for the black-antenna race of P. tharos tharos (sub- 
sequently referred to here simply as P. tharos) in Ontario, and update 
the otherwise comprehensive information available in the Ontario But- 
terfly Atlas (Holmes et al. 1991) and the annual summaries of the 
Toronto Entomological Association. 


MATERIALS AND METHODS 


Because females of P. tharos are often morphologically inseparable 
from those of P. cocyta and P. batesii, only males were used to develop 
distributional data. Males of the black-antenna race of Phyciodes tharos 
tharos were distinguished from P. cocyta males by the following charac- 
ters in order of importance: (1) unscaled portions of the tip of the an- 
tenna (nudum) being all black or mostly black (black border and lattice 
with brown steps) instead of uniformly brownish-yellow, yellow or or- 
ange; (2) antennal clubs mostly club-shaped rather than elongate; (3) 
postmedian black line on upper hindwing extending through all or most 
cubital and medial cells rather than lacking in two or more cells; (4) me- 
dian black band crossing upper forewing mostly well developed; (5) 


forewing 14-16 mm from base to apex instead of 16-18 mm; and (6) 


VOLUME 51, NUMBER 3 219 


marginal crescent patch on underside of hindwing dark brown instead 
of tan. Although a relatively small, multi-brooded race of Phycioides 
with yellow or brownish-yellow nuda, best referred to P. cocyta, occurs 
in parts of Ontario and New York state, this differs from the evidently 
more western orange antenna race of P. tharos in having the black lines 
on the upper wings less well developed. Only specimens having black 
antenna were accepted as the black-antenna race of P. tharos, but some 
_ variation was permitted in the other characters. However, the suite of 
characters associated with black antennae held together quite well. 
Black antennae were generally associated with well developed black 
lines on the upper wings for example, the most notable exception being 
a specimen at LEM from Laval (discussed below). 

Collections examined included those at Agriculture Canada in Ottawa 
(CNC), the Royal Ontario Museum in Toronto (ROM), the University 
of Guelph (UG), Lyman Entomological Museum at Ste. Anne de Belle- 
vue (LEM), and the University of Western Ontario (UWO). Members 
of the Toronto Entomological Association reporting P. tharos in recent 
season summaries (e.g., Hanks & Hess 1992, Hanks 1993, 1994, 1995) 
were contacted to confirm reports; their private collections were 
checked for other records of P. tharos. The map was produced using 
Quikmap version 2.5 (ESL Environmental Sciences Ltd., Sidney, British 
Columbia). Data for male Phyciodes tharos tharos examined in institu- 
tional collections are as follows: 


NEW YORK: Stony Point, 20 Jul 1995 (CNC); Chaumont, 2 Sep 1995 (CNC); Lock- 
port, 1 km E, 14 Sep 1995 (CNC). ONTARIO: St. Davids, 23 Aug 1930 (UWO), 1 Aug 
1932 (CNC); Brighton, 3 Aug 1932 (CNC); Fort Erie, 6 Aug 1948, 7 Aug 1948 (CNC); 
Marmora, 25 Jul 1952 (CNC); Ancaster, 26 May 1952 (CNC); Simcoe, 27 May 1953 (UG); 
Pottageville, 5 Sep 1955 (ROM); Unionville, 1956 (ROM); Don Valley, Toronto, 21 Jun 
1958, 31 May 1958 (ROM); Dunn Twp., 27 Jul 1958, 1 Aug 1958, 16 Aug 1959, 18 Aug 
1959, 16 Aug 1960, 29 Aug 1962 (UWO); Toronto, Don valley, 10 Jun 1959, 9 Aug 1959 
(ROM); Orillia, 25 Jul 1959 (ROM); Toronto, Willowdale, 5 Jun 1960 (ROM); St. Cathe- 
rines, 13 Sep 1961 (UG); Long Point, 25 May 1963 (ROM); Chaffeys Locks, 17 Jul 1963, 
22, Jul 1970, 28 May 1971, 23 May 1974 (ROM): Dunnville, 22 May 1965 (ROM); Ron- 
deau Park, 12 Jul 1965 (ROM); Wainfleet, 31 May 1969 (CNC); Oakville, 6 Jun 1976 
(UG); Dundas, 21 Jul 1981 (UG); Aberfoyle, 10 Jul 1983 (UG); Vienna, 22 Aug 1987 
(UG); 4 km W Dwyer Hill Siding, 22 Jul 1995 (CNC); Flood Rd., Rideau Twp., 22 Jul 
1995 (CNC); 1 km W Metcalfe, 22 Jul 1995 (CNC): 2 km W Metcalfe, 22 Jul 1995 (CNC); 
5 km N Metcalfe, 29 Jul 1995 (CNC); Huycks Bay, 9 Jul 1995 (CNC); Presquile Park, 15 
Jul 1995 (CNC); Bells Comers, 30 Jul 1995 (CNC); Batawa, 7 Aug 1995 (CNC); 
Kemptville, 1 Sep 1995 (CNC); Queenston, 14 Sep 1995 (CNC); Almonte, 5 km NE, 3 Sep 
1995 (CNC); Dwyer Hill Siding, 5 km SE, 4 Sep 1995 (CNC); 2.5 km SW Phragmites fen 
near Dwyer Hill Siding, 2 Sep 1995 (CNC); Mud Pond 1 km SE, 2 Sep 1995, (CNC); 1 km 
N Constance Lake, 2 Sep 1995 (CNC); Dwyer Hill Siding 3 km SE, 2 Sep 1995 (CNC); 
Prospect, Lanark 2 Sep 1995 (CNC). QUEBEC: Klock Rd., Aylmer, 2 Aug 1995 (CNC). 


Observations of oviposition were made in the field by following fe- 
males very slowly. Samples of plants upon which eggs were laid were 
identified using Semple and Heard (1987), and are deposited in the 
herbarium of Agriculture Canada in Ottawa (DAO). Larvae were reared 


220 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Phycioides tharos tharos 
(black-antenna race) 


@= males in institutional collections 
@ = males in private collections and 


reliable literature reports 


es, 78° 76° | 


Fic. 1. Distribution of the black-antenna race of Phyciodes tharos tharos in Ontario 
and adjacent regions. Based on males examined in institutional collections (dots, see Ma- 
terials and Methods) and males in private collections or reliable literature reports of males 


(half-dots). 


to adults on the same plants upon which the eggs were laid. Representa- 
tive samples of butterflies were placed in CNC. Names for vascular plants 
listed for habitats were taken largely from Gleason and Cronquist (1991). 


RESULTS AND DISCUSSION 


Distribution. Oliver (1980) suggested that the region of sympatry 
of P. tharos and P. cocyta was equivalent to Remington’s (1968) north- 
eastern suture zone and he noted a northern limit of P. tharos extending 
from central New England to southern New York, southern Ontario and 
southern Michigan. This northern limit, approximating the northern 
limit of the Carolinian or Upper Austral region of southern Ontario, was 
drawn by Scott (1986) and Opler & Malikul (1992) as extending from 
western Lake Ontario to southern Lake Huron. Figure 1 documents a 
northern limit extending farther north into the Canadian zone in the Ot- 
tawa valley north of Ottawa to 45°36’N (Lac Philippe) and to the south- 
ern edge of the Canadian Shield north of Lake Simcoe (Kirkfield and 
Orillia) at 44°38’N. 

With extensive abandoned pasture and rich alvar habitats on the 
Bruce Peninsula and Manitoulin Island, one might expect P. tharos to 


VOLUME 51, NUMBER 3 DA 


occur there, but these areas are cooler than the rest of the Ontario 
range. Phyciodes tharos was not represented among 43 Phyciodes males 
collected throughout this region at different times by J. A. Morton. It is 
also shown here for the first time in northern New York (Stony Point 
and Chaumont). 

Currently P. tharos is known in Quebec only from the Ottawa valley 
(Eardley, Lac Phillipe, Aylmer). Of 125 P. tharos (sensu lato) at LEM, 
of which approximately half are males, there is only one with black an- 
tennal clubs (Laval, A. C. Sheppard, 19 Aug 1972), but the median and 
postmedian black lines on the upper wings are poorly developed, and 
the crescent patch is tan instead of dark brown. This specimen is conse- 
quently not clearly referable to P. tharos. 

Various reports from Ontario (Hanks & Hess 1992, Hanks 1993, 1994, 
1995) from localities north of those shown on Figure 1 were found to be 
referable to P. tharos in the earlier broad sense, and those that could be 
checked proved to be P. cocyta (the only exception being a report from 
Shoal Lake Road at Hwy 17 in Kenora district (Hanks 1994) that is not 
supported by a specimen; A. Wormington, pers. comm). Phyciodes tharos 
occurs at similar latitudes in Manitoba (Klassen et al. 1989), Saskatchewan 
(Hooper 1973) and Alberta (Acorn 1993, Bird et al. 1995), and the west- 
ern race with more yellow on the nudum quite possibly occurs in at least 
the southern Kenora and Rainy River portion of Ontario. 

Status in Ontario. The relatively small number of P. tharos males 
in many institutional collections suggests the butterfly may be either 
rare in Ontario, or a recent arrival. The earliest collections of which I am 
aware are from St. Davids (43°10’N, 79°06’W) near the Niagara River 
in 1930 (UWO) and 1932 (CNC), and Brighton (44°02’N, 77°44’W) in 
Northumberland County in the eastern Lake Ontario region in 1932 
(CNC). Even at this early date, Wild (1939) noted that some authorities 
considered the darker summer brood in the Niagara frontier region to 
be a distinct variety. 

Although P. tharos sensu stricto may not be a recent arrival in the 
Lake Ontario region of Ontario, it was not noticed in the Ottawa valley 
until 1995, despite the fact that this region has received more attention 
from lepidopterists than any other part of Canada (Layberry et al. 1982). 
As soon as its habitats were understood it was found in at least 15 loca- 
tions within the district (Crolla 1996); although it proved to be quite lo- 
cal, it is locally abundant and continuously present (the earliest collected 
specimen from the district was taken in 1990 at Eardley (F. Lessard, 
pers. comm.)). Some of the places from which it currently is known are 
at or near colonies of P. batesii which were visited a number of times by 
entomologists more than 65 years ago. 

It seems unlikely that a few P. tharos would not have been taken by 


222 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the many early entomologists working in the region if it had been pres- 
ent, particularly in some of the habitats frequented by P. batesii at the 
time. Furthermore many of the current locations are abandoned pas- 
turelands, which were part of a more intensively utilized agricultural 
landscape prior to 1960. Thus, it may have moved in over the past 30 
years following abandonment of marginal pasturelands. Opler and 
Krizek (1984) speculated that P. tharos “probably expanded greatly with 
the cutting of eastern deciduous forests and expansion of agriculture.” 

On the other hand, P. tharos is quite local in the Ottawa valley and 
was previously lumped with P. cocyta, which is more conspicuous, wide- 
spread, and abundant early in the season. Consequently P. tharos may 
not have been collected “accidentally.” Appropriate species-rich habitats 
undoubtedly existed within regional alvar landscapes (see Catling & 
Brownell 1995) in pre-settlement times due to fires, and it was collected 
as far north as Marmora in 1952. 

Although P. tharos is clearly widespread in Ontario and occurs in both 
restricted natural habitats (alvars, dune slacks, prairies) and anthro- 
pogenic habitats (abandoned pastures and hay-old field complexes), the 
species does appear to be local and absent over vast areas. In contrast, 
its close relative P. cocyta is ubiquitous over much of the province in 
June and early July. Phyciodes tharos has been found in 65 locations and 
is currently known from at least 27 locations (where a location is defined 
as an area of occurrence at least 3 km from another area of occurrence). 
Consequently, its provincial ranking should be S3-S4, possibly uncom- 
mon but with sufficient occurrences to be secure, according to the Na- 
ture Conservancy of Canada system. 

Oviposition and foodplants. In southern Quebec at Aylmer (45°14'N, 
75°31’W) and in southern Ontario at Metcalfe (45°24’N, 53°75’W) and 
Dwyer Hill Siding (45°02’N, 75°49’W), females were observed oviposit- 
ing between 1300-1430 h. They flew short distances of 1-2 m, usually 
alighting and resting for 2—5 minutes, then crawled through the grasses 
and herbs 2—5 cm above the soil apparantly searching for a foodplant. 
The crawls involved distances of 0.1-0.5 m and the crawling period 
lasted for 2-30 minutes with or without resting periods of up to 10 min- 
utes (characterized by slow closing and opening of the wings to a hori- 
zontal position). During a crawl leading to oviposition a female moved 
over the same leaves 2—6 times with or without resting periods, and fi- 
nally settled on the edge of a leaf with wings slowly closing and opening 
to an angle of 45°. The abdomen was curled under the leaf and pale 
green eggs were deposited adjacent to, or on top of others, on the lower 
leaf surface. The egg clusters included 52, 58 and 64 for the three loca- 
tions listed above (respectively). Oviposition lasted approximately 20 
minutes. The plant upon which eggs were laid at each of the three sites 


VOLUME 51, NUMBER 3 WKS) 


was Aster ciliolatus Lindl. (Asteraceae), a species with elongate rhi- 
zomes that forms loose to dense patches. In all cases the females used 
relatively small, non-flowering rosette plants and placed eggs on leaves 
approximately 3 cm above the soil surface. The patches utilized were 
sparse rather than dense and lacked flowering plants, but the leaves 
upon which the eggs were laid were young and succulent. Larvae from 
the Aylmer site were reared on Aster ciliolatus from eggs to adult but- 
terflies over 68—71 days (indoors). 

Aster ciliolatus has not previously been reported as a foodplant of P. 
tharos (in either the broad or restricted sense). The range of this aster 
includes the boreal and mixed forest regions of eastern and central 
North America south to northern New England and the Great Lakes 
west across the northern edge of the prairie region (Semple & Heard 
1987). Thus, this larval foodplant is only available at the northern range 
limit of P. tharos. It was present at all sites north of Lake Ontario and 
there was a clear correlation between the amount of it and the abun- 
dance of P. tharos. Further south in Ontario Aster oolentangiensis and 
A. pilosus are suspected larval hosts. 

Habitats. The major habitat of the most northerly sites is dry, aban- 
doned pasture with short grasses and rosette-forming herbs (e.g. Dan- 
thonia spicata and Solidago nemoralis), especially where these drier ar- 
eas are interspersed with or adjacent to more moist open habitats. This 
habitat is prevalent in some alvar landscapes (i.e. areas supporting nat- 
ural limestone barrens, see Catling and Brownell 1995) north of Lake 
Ontario. Here the abandoned pastures take a long time to develop com- 
plete tree and shrub cover because of the combined effects of moisture 
extremes, periodic drought and thin soil. Abundance of Aster ciliolatus 
and overall plant diversity is lower in the sites that have experienced the 
least past disturbance. Natural alvar complexes where tree and shrub 
cover is limited by fire or cutting, as well as drought, tend to have large 
populations of both Aster ciliolatus and P. tharos as well as a high diver- 
sity of both native and introduced plant species in a mosaic of wet and 
dry habitats. Landscapes where drought is the only factor have fewer 
plant associations, many of the potential open associations having devel- 
oped tree or shrub cover. 

Reduction in overall plant diversity due to reduction in kinds of open 
habitat in the less disturbed sites reduces nectar resources for adults: 
the less extreme site conditions associated with past or present reduc- 
tion in woody cover allow flowering herbs to survive dry periods thus 
providing continuous resources for P. tharos adults. The continuity of 
nectar resources may be important to adult P. tharos since presence of 
the insect appears to be a consequence of continuous emergence (al- 
though there are definite peaks in some locations). 


224 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


In the Burnt Lands alvar complex near Almonte, Ontario, fire was his- 
torically a factor in maintaining open conditions; but with a landscape 
broken by roads and quarries and with increasing development from 
housing, fires are no longer a significant environmental factor. In one ex- 
tensive area, however, tree cover has been eliminated and shrub cover 
greatly reduced to improve the operation of radio towers, and this area 
contains a high plant diversity and P. tharos is locally abundant. The 
dominant plants in one of the major plant associations at this site include 
the grasses Agrostis stolonifera, Carex pennsylvanica, Carex umbellata, 
Danthonia spicata and Poa pratensis, and the herbs Aster ciliolatus, Co- 
mandra umbellata, Fragaria virginiana, Prunella vulgaris, Senecio pau- 
perculus, Solidago juncea, S. nemoralis, and S. ptarmicoides. 

The abandoned, rugged pastures on shallow, calcareous soil over lime- 
stone where P. tharos occurs are dominated by the grasses Agrostis 
stolonifera, Danthonia spicata, Dactylis glomerata, Festuca spp., Poa 
compressa, Panicum philadelphicum, Sporobolus vaginiflorus, and the 
herbs Aster ciliolatus, Daucus carota, Echium vulgare, Fragaria virgini- 
ana, Hieracium pilloselloides, Leucanthemum vulgare, Melilotus alba, 
Origamum vulgare, Prunella vulgaris, Rudbeckia hirta, Senecio pauper- 
culus, Solidago canadensis, S. nemoralis, Trifolium spp. and Vicia 
cracca. The continuity of blossoms from spring through summer to fall 
in these habitats is to a large extent a result of the presence of intro- 
duced species. The nearly complete restriction of P. tharos to the dry al- 
var and abandoned limestone pasture habitats at its northern range limit 
could relate to the relatively warmer microclimate of these dry, open 
sites as well as to both adult and larval foodplant availability. 

Farther south of the alvar landscapes and particularly in the Carolin- 
ian region of Ontario and northern New York, the main species of Aster 
associated with P. tharos at several sites is the white A. pilosus. Adult P. 
tharos occur in association with this species in natural habitats such as 
dune slacks dominated by the graminoids (grasses and sedges) Cladium 
mariscoides, Poa compressa, and Schizachyrium scoparium, and in old 
fields dominated by the grasses Agrostis spp., Dactylis glomerata, 
Phleum pratense, Poa compressa, Poa pratensis, and the herbs Daucus 
carota, Solidago canadensis, and S. nemoralis. Occupied habitats are 
primarily in areas of sandy soils subject to moisture extremes and where 
encroachment of shrubs such as Cornus racemosa is relatively pro- 
longed. It also occupies hayfields that are cut once, but in these habitats 
there is probably an enforced displacement after cutting to adjacent un- 
cut or abandoned fields. Cutting of hay results in a second or late 
blooming of Daucus carota, Trifolium spp., and other species, thus im- 
proving the local continuity of adult food resources. The hayfields often 
have Trifolium spp., Lotus corniculatus, Dipsacus sylvestris, Ambrosia 


VOLUME 51, NUMBER 3 225 


artemisiifolia among the dominants, the only prominent aster being 
Aster pilosus var. pilosus. In the dune slacks the prominent aster is usu- 
ally Aster pilosus var. pringlei, but A. dumosus is also present. 

In one prairie situation in southern Ontario (Brant County), P. tharos 
occurred in a hilly area dominated by Schizachyrium scoparium, 
Sorghastrum nutans and Aster oolentangiensis with no other Aster spe- 
cies evident. This site had been impacted previously by grazing and had 
many alien species. Other southern Ontario prairies surveyed were 
without populations of P. tharos. 

In many cases P. tharos was found to be absent from areas within the 
general region of occurrence indicated in Figure 1. Some of the sand 
barrens and granite rock barrens searched had large populations of 
Aster ciliolatus, but there were periods of 4 weeks or more during the 
summer when nectar resources were essentially adsent. In contrast, the 
closely related, and more or less univoltine, P. cocyta was present in 
most if not all of these sites. All sites where P. tharos was found includ- 
ing abandoned pastures, disturbed alvar, dune complexes, and hayfield- 
old field landscapes, had in common a high floristic diversity and a con- 
tinuous supply of adult food resources, as a result of a mosaic of 
different habitats lacking woody cover. 

Flight period. The earliest adult flight dates for Ontario are 22—26 
May (Dunnville to the Ottawa district, respectively). The latest dates 
range from 13 October at Ottawa (pers. obs.) to 23 October on Point 
Pelee (Hanks & Hess 1992, reported by A. Wormington). Adult emer- 
gence appears to be almost continuous, with worm and fresh specimens 
of both males and females being found in most samples. At locations 
where there are large populations, adults were encountered from early 
June to early October. Appearance of the butterflies seems to depend as 
much on the weather as on a regular schedule of consecutive broods, 
but there were three clear peaks of abundance in 1995 (early June, mid- 
July to early August, and late August to early September). 


ACKNOWLEDGMENTS 


L. F. Gall of Yale University, A. Hanks of Aurora, Ontario, D. Lafontaine of Agriculture 
Canada in Ottawa, P. Opler of Fort Collins, Colorado and J. Scott of Lakewood, Colorado, 
all provided helpful comments on the manuscript. A. Wormington of Leamington, On- 
tario, and J. Scott provided information on the characteristics of P. tharos in their respec- 
tive regions. J. Morton of Waterloo, Ontario kindly made his collection from Manitoulin 
Island available for study. Many entomologists including members of the Toronto Ento- 
mological Association provided and/or corrected records: R. Bowles (Orillia), J. Crolla (Ot- 
tawa), S. Daniels (Toronto), N. Escott (Thunder Bay), N. Ironside (Orillia), R. Layberry 
(Ottawa), F. Lessard (Deux Montagnes), W. McIlveen (Acton), L. Taman (Matachewan). 


LITERATURE CITED 


ACORN, J. 1993. Butterflies of Alberta. Lone Pine Publishing, Edmonton, Alberta. 141 pp. 
BIRD, C. D., G. J. HILCHIE, N. G. KONDLA, E. M. PIKE & F. A. H. SPERLING. 1995. Al- 
berta butterflies. Provincial Museum of Alberta, Edmonton. 347 pp. 


226 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


CATLING, P. M. & V. R. BROWNELL. 1995. A review of the alvars of the Great Lakes re- 
ion: distribution, floristic composition, biogeography and protection. Can. Field Nat. 
109(2):143=172- 

CROLLA, J. 1996. The Pearl Crescent (Phyciodes tharos) in the Ottawa District. Trail & 
Landscape 30(2):51—57. 

GLEASON, H.A. & A. CRONQUIST. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada. Second Edition. The New York Botanical Garden, Bronx, 
New York. 910 pp. 

HANKS, A. J. 1993. Butterflies of Ontario and summaries of Lepidoptera encountered in 
Ontario in 1992. Toronto Entomol. Assoc. Occ. Publ. 25—93. 102 pp. 

. 1994. Butterflies of Ontario and summaries of Lepidoptera encountered in On- 

tario in 1993. Toronto Entomol. Assoc. Occ. Publ. 26—94. 100 pp. 

. 1995. Butterflies of Ontario and summaries of Lepidoptera encountered in On- 
tario in 1994. Toronto Entomol. Assoc. Occ. Publ. 27—95. 92 pp. 

HANKS, A. J. & Q. F. HEss. 1992. Butterflies of Ontario and summaries of Lepidoptera en- 
countered in Ontario in 1991. Toronto Entomol. Assoc. Occ. Publ. 24—92. 94 pp. 
HOLMES, A. M., Q. F. HEss, R. R. TASKER AND A. J. HANKS. 1991. The Ontario Butterfly 

Atlas. Toronto Entomol. Assoc. 167 pp. 

Hooper, R. R. 1973. The butterflies of Saskatchewan. Museum of Natural History, 
Regina. 216 pp. 

KLASSEN, P., A. R. WESTWOOD, W. B. PRESTON & W. B. MCKILLOP. 1989. The butterflies 
of Manitoba. Manitoba Museum of Man and Nature, Winnipeg. 290 pp. 

LAYBERRY, R. A., J. D. LAFONTAINE & P. W. HALL. 1982. Butterflies of the Ottawa district. 
Trail and Landscape 16(1):3—59. 

OLIVER, C. G. 1980. Phenotypic differentiation and hybrid breakdown within Phyciodes 
“tharos” (Lepidoptera: Nymphalidae) in the northeastern United States. Ann. Ento- 
mol. Soc. Am. 73:715—721. 

OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins 
Univ. Press, Baltimore. 294 pp. 

OPLER, P. A. AND V. MALIKUL. 1992. A field guide to the butterflies. Houghton Mifflin 
Co., New York. 396 pp. 

ScorTT, J. A. 1986a. The butterflies of North America, a natural history and field guide. 
Stanford University Press, Stanford, California. 583 pp. 

. 1986b. The courtship of Phyciodes, and the relationship between Phyciodes 

tharos tharos and Phyciodes tharos morpheus (=pascoensis) in Colorado. Papilio, new 

series, 5:1—8. 

. 1994. Biology and systematics of Phyciodes. Papilio, new series, 7:1—120. 

SEMPLE, J. C. & S. B. HEARD. 1987. The asters of Ontario: Aster L. and Virgulus Raf. 
(Compsitae: Astereae). Univ. Waterloo (Waterloo, Ontario), Biol. Ser. No. 30. 88 pp. 

WILD, W. 1939. The butterflies of the Niagara frontier region. Buffalo Soc. Natural Sci. 
Bull. 19(1):1—55. 


Received for publication 20 October 1995; revised and accepted 13 May 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 227-236 


TWO NEW SPECIES OF ASTERACEAE-FEEDING 
BUCCULATRIX (BUCCULATRICIDAE) FROM CALIFORNIA 


DANIEL Z. RUBINOFF 


Department of Environmental Science Policy and Management, 
University of California, Berkeley, California 94720, USA 


AND 


KENDALL H. OSBORNE 
Department of Entomology, University of California, Riverside, California 92521, USA 


ABSTRACT. Bucculatrix tetradymiae, new species and Bucculatrix dominatrix, 
new species are described and illustrated. Bucculatrix tetradymiae feeds on Tetradymia 
axillaris (Strother) (Asteraceae) and Bucculatrix dominatrix feeds on Baccharis pilularis 
(de Candolle) (Asteraceae) and can be distinguished from sympatric, Baccharis-feeding 
Bucculatrix variabilis (Braun) and Bucculatrix separabilis (Braun) by its larger size, dis- 
tinct forewing pattern, and genitalia. 


Additional key words: Tetrydamia, Baccharis, leaf miner, Lyonetiidae, microlepi- 
doptera. 


Zimmerman (1978) resurrected the family Bucculatricidae (Bucculat- 
rigidae) from Lyonetiidae, a move first proposed by Fracker (1915). The 
family is easily discerned, being characterized by an “elongate pointed 
face, tufted head, basal eye-cap of the antenna and, in the male, the 
notched first segment of the flagellum . . .” (Braun 1963). The larvae 
typically are leaf miners in the early instars, and then become external 
feeders, although a few species mature in the mine and some are gall- 
makers. For a complete description of the family refer to Braun (1963). 
The family is cosmopolitan with 222 species described from all land 
forms except New Zealand (Heppner 1991). More than 100 occur in 
North America, mostly in arid regions (Braun 1963). Larvae of many 
western species feed on Asteraceae, including the two species we de- 
scribe here. 

On 12 April 1993 in the western margin of the Mojave desert, Califor- 
nia, we collected cocoons affixed to stems of the spiny shrub Tetradymia 
axillaris. Cut branches with attached cocoons were immersed in buckets 
of water and transported to our homes in central California and so main- 
tained. Adult Bucculatrix emerged during late April and early May. 

David Wagner, John De Benedictis and Jerry Powell collected a new 
Bucculatrix on Mt. San Bruno (San Mateo Co.), where it is sympatric 
with the Baccharis-feeding Bucculatrix variabilis and Bucculatrix sepa- 
rabilis; De Benedictis et al. (1990:p.20) briefly described the adult co- 
coon and larval biology of this species without naming it. This species 
has also been reared from Marin County and Sonoma County by Jerry 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 1-4. 1, Bucculatrix tetradymiae paratype female. California: San Bernardino 
Co.: Oro Grande Wash, 5 mi. W. of Hesperia, 12 April 1993 (Rubinoff and Osborne), 
reared from cocoon on Tetradymia axillaris, ex pupa 20 April 1993 (JAP 93D45). 2, B 
dominatrix type female. California: Marin Co.: Ring Mountain, 19 April 1991 (J. A. Pow- 
ell), reared from cocoon on Baccharis pilularis, ex pupa 1-2 May 1991 (JAP 91D 16). 3, 
Bucculatrix variabilis. California: Monterey Co. Big Creek Reserve (UCNRS), 1-3 May 
1992 (J. A. Powell) reared from cocoon on Baccharis pilularis, ex pupa 6 May 1992 (JAP 
92E11). 4, Bucculatrix separabilis. California: Napa Co. 1 mile SE of Angin 20 May 
1980 (J. A. Powell). 


Powell and David Wagner, respectively. Both new species were sexed on 
the basis of frenulum morphology and external genitalia. 


Bucculatrix tetradymiae Osborne and Rubinoff, new species 
(Figs. 1, 5-8) 


Description. Head. White; antenna white with white basal eye-cap and dark brown 
annulations. Thorax. White. Forewing. Length: mean 3.4mm (range 3.2—3.8mm, n = 29). 
Dorsal surface lustrous white with distal half dominated by roughly equal-sized postmedial 
and submarginal blotches partially separated by white at anal angle and costa. Blotches, 
ochreous brown, tipped with dark brown. Postmedial blotch often bisected by diffuse lon- 
gitudinal white or light brown. Occasionally blotches greatly reduced. Below fold, in prox- 
imal extreme of postmedial blotch, a very dark patch of completely and partially dark 
brown scales. Series of white, dark-tipped scales running dorsally from apex along margin 
to merge with distal edge of dark field. Usually a small patch of several dark brown tipped 
scales on costa at 1/3 from base to apex. Cilia brown above brown field on costa, white sub- 
apically, brown in streak at apex on apical end of marginal chain of brown scales, all white 
along margin. Ventral surface of forewing gray-brown. Hindwing. Gray. dorsally, gray- 
brown ventrally with gray cilia. Leg. White with dark brown annulations on distal ends of 
tarsal segments. Metathoracic tibiae with long white cilia. Abdomen. Gray. Terminal scales 
elongate and dark gray dorsally in females. 

Male genitalia (Figs. 5, 6 drawn from YFH prep no. 0911). Valva elongate, slightly 
curved medially with fine distal setation. Socii divergent, elongate, with long setae on dis- 


VOLUME 51, NUMBER 3 229 


Fics. 5,6. 5, B. tetradymiae paratype male genitalia. 6, aedeagus of same. Slide YFH 
0911. California: San Bernardino Co., Oro Grande Wash, 5 mi. W. of Hesperia, 25 April 
1993 (D. Rubinoff). Scale bars = 1 mm. 


230 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Vane) 


Fic. 7. B. tetradymiae paratype female genitalia. Slide YFH 0905. California: San 
Bernardino Co., Oro Grande Wash, 5 mi. W. of Hesperia, 25 April 1993 (D. Rubinoff ). 


Scale bars = 1 mm. 


VOLUME 51, NUMBER 3 Zoi 


Fic. 8. B. tetradymiae paratype female genitalia, corpus bursae. Slide YFH 0906. Cal- 
ifornia: San Bernardino Co., Oro Grande Wash, 5 mi. W. of Hesperia, 25 April 1993 (D. 
Rubinoff ). Scale bars = 1 mm. 


tal third. Gnathos absent. Vinculum narrow, well sclerotized. Aedeagus straight, gradually 
tapering to slender apex. 

Female genitalia (Fig. 7 drawn from YFH prep no. 0905, Fig. 8 drawn from YFH 
prep no. 0906). Ductus bursae not sclerotized. Margins of ostium bursae weakly sclero- 
tized. Ostium in anterior margin of abdominal segment eight. Ninth tergum with trans- 
verse row of long, stout setae at mid-segment. Posterior apophyses about length of ab- 
dominal segment eight, well sclerotized. Anterior apophyses absent. Ductus seminalis 
arising on corpus bursae near base of ductus bursae. Signum strong, nearly encircling pos- 
terior half of bursa, ribbed with long aciculae; converging toward base of ductus bursae. 

Type specimens. Holotype 3, California: San Bernardino Co.: Oro Grande Wash, 5 
mi. W. of Hesperia, 12 April 1993 (Rubinoff and Osborne), reared from cocoon on 
Tetradymia axillaris, ex pupa 20 April 1993. Paratypes (n = 49): California: San Bernardino 
Co.: Oro Grande Wash, 5 mi. W. of Hesperia, 1 5, 1 2, 12 April 1993 (Rubinoff and Os- 
borne), reared from cocoons on Tetradymia axillaris, ex pupa 20 April 1993 ; also 2 4, 6 2, 
25 April 1993 (Rubinoff and Osborne) reared from cocoons on Tetradymia axillaris, ex 
pupa 1—3 May 1993 (JAP 93D48); also 17 4, 12, 27 April 1996 (Osborne); also 18 4, 3 2, 29 
April 1996 (Osborne). The holotype and 26 paratypes are deposited at the Essig Museum 
of Entomology, University of California, Berkeley (UCB); 4 paratypes are deposited in the 
U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C. 


232 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(USNM); 4 paratypes are deposited in the University of Connecticut collection at Storrs 
(UCONN); 3 paratypes are deposited in the Los Angeles County Museum (LACM); 3 
paratypes are deposited in the Canadian National Collection of Insects (CNCI); 3 paratypes 
are deposited in the California Academy of Sciences (CASC); 4 paratypes remain in the col- 
lection of Kendall Osbome and 2 paratypes in the collection of Daniel Rubinoff. 

Diagnosis and discussion. Bucculatrix tetradymiae is a small, white bucculatricid 
with brown blotches dominating the outer half of the forewing. B. tetrydamiae keys out to 
Bucculatrix packardella (Chambers) (of the eastern United States) in Braun’s (1963) key 
to the adults based on maculation. The oak-feeding B. packardella differs from B. 
tetradymiae by the brown speckles over the face, eye-caps and thorax, and by the dusting 
of brown-tipped scales over the forewing base. On the basis of genitalic characters and 
host specialization, B. tetradymiae falls within Braun’s (1963) section II of Bucculatrix 
which contains the majority of North American, Asteraceae-feeding Bucculatrix; B. 
packardella is assigned to section IV. 

Adult moths are found in close association with the host, Tetradymia axillaris, in the 
western Mojave Desert. The early stages probably mine the glabrose, fascicular leaves 
while the late stage larvae feed externally. The white cocoons were fixed lengthwise usu- 
ally against axillary spines of the host, are mostly smooth with slight ridges discernible on 
the caudal terminus. This unusual smoothness (most Bucculatrix cocoons have conspicu- 
ous sculpturing) is not due to the cocoon-smoothing effects of parasitoids, as hypothesized 
by Braun (1963) since many of our cocoons were viable. Adults emerged between 20 April 
and 10 May 1994 in captivity from cocoons kept both indoors (room temperature) and 
outdoors in Berkeley, California. 

We saw many adults on Tetradymia at Oro Grande Wash on 23 April 1994, but we 
found none when we returned on 13 May 1994. Adults are active from at least 1600 to 
2000 hours. Females extrude the papillae anales while resting on the host plant and males 
rapidly crawl (about 1 cm per second) along the length of stems usually searching each ax- 
illary spine in sequence along a stem, then crawling or sometimes flying to a new stem. An 
observed mating lasted 45 minutes. All known specimens come from the type locality al- 
though the species should be expected over much of the Mojave desert and may extend 
over the range of its host. 


Bucculatrix dominatrix Rubinoff and Osborne, new species 
(Figs. 2, 9-12) 


Description. Head. Grayish white, tuft brown and white hairs, eye-caps gray; antenna 
black and white banded. Thorax. grayish-white with scattered brown-tipped scales. 
Forewing. Length: mean: 5.5 mm (range: 4.0—6.lmm, n = 32), mottled brown, prominent 
longitudinal white streak extending above Cu from base along the discal cell, then abruptly 
jutting at acute angle towards costal margin. Just below Cu at acute angle in white streak, 
small, very dark patch of scales, just below (fold) vein. Distally, mottling becoming lighter, 
ending nearly white and surrounded by fuscous-tipped apical scales. Cilia gray. Ventral sur- 
face iridescent brown. Hindwing. Pale gray, ventral surface iridescent brown. Leg. Gray with 
distal end of tibia turning black. Tarsi banded with black and white. Abdomen. Silvery gray. 

Male genitalia (Figs. 9, 10 drawn from YFH prep no. 0903). Valva elongate, point- 
ing inwards at the setose distal ends. Soci divergent, very broad, more setose apically, and 
fused with tegumen. Saccus absent. Vinculum well-sclerotized band. Uncus poorly sclero- 
tized with distal end obtuse. Aedeagus arising from tapered annellus. Distinguished from 
male genitalia of B. variabilis by much shorter, stout socii; very similar to male genitalia of 
B. separabilis. 

Female genitalia (Figs. 11, 12 drawn from YFH prep no. 0907). Ostium in deep, 
well-sclerotized, cup-shaped chamber. Corpus bursae ovoid. Ductus seminalis originating 
on corpus bursae just dorsal to junction of ductus bursae and corpus bursae, no expansion 
of ductus seminalis apparent. Signum narrow, forming densely spined band of ribs at the 
posterior end of corpus bursae just anterior of where ductus seminalis and ductus bursae 
join corpus bursae. 

Type specimens. Holotype °, California: Marin Co.: Ring Mountain, 19 April 1991 


VOLUME 51, NUMBER 3 238 


Fics. 9,10. 9, Paratype male genitalia of B. dominatrix. 10, aedeagus of same. Slide 
YFH 0903. California: Marin Co., Ring Mt., 19 April 1991 (J. A. Powell) reared from co- 
coon on Baccharis pilularis, ex pupa 1-5 May 1991 (JAP 91D16). Scale bars = 1 mm. 


(J. A. Powell), reared from cocoon on Baccharis pilularis, ex pupa 1-2 May 1991 (JAP 
91D16). Paratypes (n = 42): California: Marin Co.: Ring Mountain, 6 d, 5 2, 11 April 1994 
(J. A. Powell), reared from cocoons on Baccharis pilularis, ex pupa 20-26 April 1994 (JAP 
94D53); also, 6 5, 22 19 April 1991 (J. A. Powell), reared from cocoons on Baccharis pilu- 
laris, ex pupa 1-5 May 1991 (JAP 91D16); San Mateo Co.: Mt. San Bruno County Park, 3 
d, 2°, 19 April 1988 (J. A. DeBenedictis) at b.]., also 3 d, 4 2, 21 April 1983 (J. A. DeBene- 
dictis), reared from cocoons on Baccharis pilularis, ex pupa 11-17 May 1983 (JADeB 
83111-E), also 1 2°, 14 April 1983 (J. B. Whitfield), reared from cocoon on Baccharis pilu- 
laris, ex pupa 3 May 1983 (JAP 83D70); Alameda Co.: Strawberry Canyon, UCB campus, 
1 5, b.1. trap, 1 July 1991 (J. A. Powell), also 1 3, 20 June 1990 (J. A. Powell); Sonoma Co.: 
1 mi. SE Bodega Bay, 4 4, 4 2, 20 April 1983 (D. L. Wagner), reared from cocoons on Bac- 
charis pilularis, ex pupa 3-24 May 1983 (JAP 83D110). The holotype and 20 paratypes 
are deposited in the Essig Museum of Entomology at the University of California, Berke- 
ley (UCB); 3 paratypes are deposited in the U.S. National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. (USNM); 11 paratypes are deposited in the 
University of Connecticut collection at Storrs (UCONN); 2 paratypes are deposited in the 
Canadian National Collection of Insects (CNCI); 2 paratypes are deposited in the Los An- 
geles County Museum (LACM); 2 paratypes are deposited in the California Academy of 
Sciences (CASC); | paratype remains in the collection of Kendall Osborne and 1 paratype 
in the collection of Daniel Rubinoff. 

Diagnosis and discussion. Bucculatrix dominatrix is relatively large and distin- 
guishable by a prominent white streak on the upper part of the discal cell, extending two- 


234 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


{\ ie / 


Fic. 11. Paratype female genitalia of B. dominatrix. Slide YFH 0907. California: 


Marin Co. Ring Mt., 19 April 1991 (J. A. Powell) reared from cocoon on Baccharis pilu- 
laris, ex pupa 1-5 May 1991 (JAP 91D 16). Scale bars = 1 mm. 


VOLUME 51, NUMBER 3 235 


i)’ 
u Ag y 
rey: age 


Fic. 12. Paratype female genitalia of B. dominatrix, bursa copulatrix. Slide YFH 0907. 
California: Marin Co. Ring Mt., 19 April 1991 (J. A. Powell) reared from cocoon on Bac- 
charis pilularis, ex pupa 1-5 May 1991 (JAP 91D 16). Scale bars = 1 mm. 


236 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


thirds the length of the wing. B. dominatrix is probably a close relative of both Bucculatrix 
variabilis (Fig. 3) and B. separabilis (Fig. 4) and phenotypically resembles the former. The 
new species is 1.5 to 2 times larger than B. variabilis. The forewing pattern is diagnostic: 
B. variabilis has two transverse white bands whereas B. dominatrix has a single, longitudi- 
nal white streak. There is also a tuft of dark scales below the longitudinal band that is not 
present in B. variabilis. B. dominatrix is larger and much darker than B. separabilis but 
the genitalia are nearly identical, indicating a probable sister species relationship between 
the taxa. Phenotypic and genitalic similarities to the two aforementioned species merit its 
placement in Section I, subsection A, of Braun (1963). Average cocoon length is 8.1 mm 
(range: 6.5-11.5 mm, n = 20). They are strongly ribbed and pinkish when occupied 
(DeBenedictis et al. 1990) turning white after emergence. 

Bucculatrix dominatrix may be widely distributed in coastal central California, where 
the hostplant is found. It feeds on Baccharis pilularis and can be found feeding on the 
same plants as both Bucculatrix variabilis and B. separabilis. 


ACKNOWLEDGMENTS 


We thank Jerry Powell for drawing our attention to the novelty of B. tetradymiae and 
the need for description of B. dominatrix, and his encouragement and tireless consulta- 
tion during our efforts, Yu-Feng Hsu for preparing the genitalia slides and assistance with 
the manuscript, David Wagner for loaning specimens and, along with John De Benedictis, 
reviewing our manuscript in addition to collecting some of the first B. dominatrix. Addi- 
tional thanks go to Don Davis for confirming the novelty of B. tetradymiae, Michelle Mar- 
vier for confirming our identification of the Tetradymia host, and to Larry Serpa of The 
Nature Conservancy, Tiburon, California for access to the Ring Mountain Preserve in 
Marin County. Comments from Lawrence Gall and two anonymous reviewers improved 
the quality of this paper. We are grateful to Sean O'Keefe for his skillful preparation of the 
photographs. Drawings were done by K. H. Osborne. Both authors contributed equally to 
the manuscript. 


LITERATURE CITED 


BRAUN, A. F. 1963. The genus Bucculatrix in America north of Mexico (Microlepi- 
doptera). Mem. Am. Entomol. Soc. 18. 

DE BENEDICTIS, J. A., D. L. WAGNER & J. B. WHITFIELD. 1990. Larval hosts of mi- 
crolepidoptera of the San Bruno Mountains, California. Atala 16:14—35. 

FRACKER, S. B. 1915. The classification of lepidopterous larvae. Illinois Biol. Monographs 
1:1-169. 

HEPPNER, J. B. 1991. Faunal regions and the diversity of Lepidoptera. Tropical Lepi- 
doptera 2 (suppl. 1):1—85. 

ZIMMERMAN, E. C. 1978. Insects of Hawaii, p. 719, v. 9, Microlepidoptera. Part I. The 
University Press of Hawaii, Honolulu. 


Received for publication 13 November 1995; revised and accepted 13 July 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 237-248 


A REVISION OF THE CERASTIS CORNUTA GROUP OF THE 
GENUS CERASTIS SUBGENUS METALEPSIS (NOCTUIDAE) 


LARS CRABO 


Thomas Burke Memorial Washington State Museum, Seattle, 
Washington 98225, USA 


AND 


J. DONALD LAFONTAINE 


Biological Resources Division, Centre for Land and 
Biological Resources Research, Agriculture and Agri-Food Canada, 
Ottawa K1A 0C6, Ontario, Canada 


ABSTRACT. The noctuid genera Cerastis Ochsenheimer and Metalepsis Grote are 
reviewed resulting in revision of Metalepsis to a subgenus of Cerastis. The C. cornuta 
group of Cerastis subgenus Metalepsis is defined. In addition to cornuta Grote, this group 
contains three new species: C. enigmatica, new species from the Pacific Northwest; C. 
robertsoni, new species from southern California; and C. gloriosa, new species from the 
west coast of the continental United States. The species are illustrated and a key for their 
identification is presented. 


Additional key words: California; Pacific Northwest; sphagnum bog; coastal 
chapparal. 


Cerastis Ochsenheimer is a genus of 13 species of medium-sized 
moths which occur in temperate forests of the Holarctic region. They 
are unusual in the subfamily Noctuinae in that the adults are active in 
early spring, flying with species in the subfamilies Psaphidinae, Hadeni- 
nae and some Ipimorphinae (Xylenini: Lithophane, Eupsilia). 

The North American species of Cerastis were previously arranged in 
two genera. Three species with foretibial spines constituted Metalepsis 
Grote. The type species of Metalepsis, cornuta Grote, 1874, occurs on 
the west coast and was thought to have a range extending north from its 
type locality, California, to the Alaska Panhandle. A new species allied to 
C. cornuta was recently discovered on the coast of California and Wash- 
ington. Subsequent study of the populations previously considered to be 
C. cornuta revealed that these consisted of three allopatric species. 
Cerastis cornuta is restricted to central California and populations to the 
north and south of its range, although superficially similar, are distinct 
species differentiated by genitalic characters. 

The relationship of Metalepsis to Cerastis was re-evaluated as part of 
our study of the C. cornuta group. McDunnough (1927), recognizing 
the close relationship of these genera, stated that “the male genitalia of 
the two genotypes are practically identical.” He retained Metalepsis for 
the species with sclerotized setae (“spines”) on the tibia of the protho- 
racic leg. This treatment was followed in subsequent checklists and cat- 


238 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


alogs of North American noctuid moths Mees ateeCIEs 1938, Fran- 
clemont & Todd 1983, Poole 1989). 

We retain Metalepsis as a subgenus, new status, of Cerastis. These 
two subgenera are distinguished by the shape of the digitus, the pres- 
ence or absence of sclerotized foretibial setae, and the shape of segment 
A8 in the female. In subgenus Cerastis the digitus is immediately distal 
to the clasper and is free from the inner surface of the valve for most of 
its length, the foretibiae are devoid of setae, and abdominal segment 
eight of the female forms lobes which project into each side of the os- 
tium bursae. In subgenus Metalepsis the digitus is fused to the inner 
surface of the valve for most of its length with only the apical third to 
quarter free, the tibia of the first leg bears complete inner and partial 
outer rows of sclerotized setae, and the ostium bursae is without lobes. 
There are seven species in subgenus Cerastis, tenebrifera (Walker 1865) 
in North America and six in Eurasia (Fibiger 1993). The six species in 
subgenus Metalepsis are restricted to North America. The revised status 
of Metalepsis results in the following new combinations for these spe- 
cies: Cerastis cornuta, new combination; Cerastis fishii, new combi- 
nation; and Cerastis salicarum (Walker 1857), new combination. 

The species in these subgenera share derived characters of adults and 
larvae indicating that they form a monophyletic group and should be 
united in one genus: male antennae bipectinate (reduced in C. fishii 
(Grote 1878) and the Old World C. rubricosa |Denis and Schiffer- 
miiller] 1775); head and thorax covered with hairlike scales; male geni- 
talia with anellus sclerotized laterally and covered with short stout 
spines, valve with digitus but lacking a corona, and vesica coiled in a sin- 
gle loop with one or more large basal cornuti; female genitalia with 
bisaccate bursa and membranous ventral cleft in the sclerotization of the 
ductus bursae; larva with ridges on inner surface of mandible extending 
to the cutting margin without a tooth on the inner surface, and with 
stemmata 3 and 4 very close together, almost touching. 

The presence of the digitus is unusual in the subfamily Noctuinae and 
is probably a primitive condition. Choephora Grote, type species Choe- 
phora fungorum Grote & Robinson, 1868, is the genus most closely re- 
lated to Cerastis. The two genera share the distinctive characters of the 
anellus and larvae and the antennae, vestiture and shape of the valves 
are similar. Choephora differs from Cerastis in lacking the digitus on the 
valve and in its mid-summer flight season. 

The Cerastis cornuta group includes four species, three of which are 
described as new. All four are restricted to the west coast of North 
America. The adults are easily distinguished from other species in the 
subgenus by the distinctive forewing spots which are conspicuously out- 
lined with pale scales. The elongate oblique orbicular and reniform 


VOLUME 51, NUMBER 3 239 


spots are fused across the median space in most specimens, forming a 
broad V. The uncus of the male genitalia is broadest apically in the C. 
cornuta group but subapically in other Cerastis. The four species are su- 
perficially similar and can most reliably be distinguished by genitalic 
characters, however, subtle differences in wing pattern and length can 
be used to separate some specimens without dissection. Identification is 
simplified since only Cerastis gloriosa occurs sympatrically with other 
species in the group. Cerastis gloriosa is the most distinctive species and 
can usually be identified without dissection, allowing the three remain- 
ing species to be tentatively identified by geographic location. 

The terminology for wing pattern and genitalia structures follows that 
used in the Moths of America North of Mexico series. These are illus- 
trated by Hodges (1971: vi—vii). 


Key to adults of the cornuta group of Cerastis 


1. Orbicular spot mostly filled with white or pale yellow (Figs. 1-2); larger species, 

forewing length 14-17 mm; male antenna 2x as wide as the central shaft; 

clasper short, projecting posteriorly, not reaching dorsal margin of valve (Fig. 9); 

appendix bursae smaller than corpus bursae with ductus seminalis at posterior 

aime! (Ita, 1S) aie glee ie: ict lapnte ana ae Simi eral al craic arar Pete eI Ets Fane ae C. gloriosa 
1’. Orbicular spot gray or brown, usually outlined in yellow (Figs. 3—8); smaller 

species, forewing length 11-15 mm; male antenna 3x as wide as central shaft; 

clasper extending posterodorsally beyond dorsal margin of valve (Figs. 10-12); 

appendix bursae larger than corpus bursae with ductus seminalis at anterior end 

ee le eS, Oi ee le ee Nees ee 2 
2. Vesica with a basal coil, bent ventrad 90° near base then to left, curving in an 

arc over 180° to project posterolaterally to the right (Fig. 10); appendix bursae 

joined to corpus bursae 1/3 from posterior end and extending obliquely towards 

corpus bursae anteriorly (Fig. 14); occurring in Cascades and West Coast from 

the Alaska Panhandle to northern California (Humboldt County) .... C. enigmatica 
2’, Vesica bent 90° once or twice at base, then nearly straight (Figs. 11-12); appen- 

dix bursae joined to corpus bursae at posterior end and extending obliquely away 

from corpus bursae anteriorly (Figs. 15-16); occurring in western California ..... 
3. Vesica bending 90° ventrad near base, then straight (Fig. 11); appendix bursae 

even in width from base to apex, nearly straight, projecting slightly anteriorly at 

apex (Fig. 15); ductus bursae shorter, 1.5x as long as wide; occurring along the 

coast from San Francisco Bay torSonomas: Counties yey es 2 te C. cornuta 
3°. Vesica with two 90° bends near base, bending ventrad for distance equal to 

aedaeagus width and then laterally to left (Fig. 12); appendix bursae constricted 

apically, curving dorsally at apex (Fig. 16); ductus bursae 2x as long as wide; 

occuring in Coast Range from Monterey Bay to Santa Barbara County C. robertsoni 


Cerastis gloriosa Crabo & Lafontaine, new species 
(eters. IL, 4, Gl, 183) 


Description. Male (Figs. 1, 2): Head: dark reddish brown; palpi concolorous. Eyes: 
black. Antennae: bipectinate, 2x as wide as central shaft. Thorax: dark reddish brown, col- 
lar lighter, reddish brown with darker lines. Foretibiae: spined. Forewing: length 14-17 
mm; dark reddish brown, suffused with variable amounts of black in discal cell; basal area 
with gray scales and violaceous tint; terminal area lighter than remainder of forewing, also 
with violaceous tint; lines double, black, pale filled; basal line sinuous; antemedial line 
oblique, gently excurved, notched sharply basad in fold; median shade usually obsolete, 
when present weak, wavy, evident only from reniform spot to inner margin; postmedial 


240 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-8. Adults of Cerastis spp. 1, C. gloriosa Crabo & Laf., paratype d, North Bay Bog, 
Grays Harbor County, Washington; 2, C. gloriosa Crabo & Laf., 3, Mill Valley, California; 3, 
C. enigmatica Laf. & Crabo, holotype ¢, Vancouver, British Columbia; 4, C. enigmatica Laf. 
& Crabo, paratype °, Wellington, British Columbia; 5, C. cornuta (Grote), 3, Inverness, 
Marin County, California; 6, C. cornuta (Grote), 3, Bodega, Sonoma County, California; 7, C. 
robertsoni Laf. & Crabo, paratype ¢, Gaviota Pass, Santa Barbara County, California; 8, C. 
robertsoni Laf. & Crabo, paratype °, Gaviota Pass, Santa Barbara County, California. 


VOLUME 51, NUMBER 3 QA] 


12 


Fics. 9-12. Male genitalia of Cerastis spp. 9, C. gloriosa Crabo & Laf., Mill Valley, 
California, CNC 10296; 10, C. enigmatica Laf. & Crabo, Ketchikan, Alaska, CNC 11140; 
11, C. cornuta (Grt.), Oakland, California, CNC 11139; 12, C. robertsoni Laf. & Crabo, 
Gaviota Pass, Santa Barbara County, California, CNC 11135. 


line dentate, lower half nearly straight, excurved opposite cell; subterminal line weak, pale, 
undulating, with strong wedge-shaped black patches proximally opposite discal cell be- 
tween median veins and near costa on radial veins; terminal line thin, weakly scalloped; 
claviform spot narrow, black; orbicular spot large, an elongate ellipse, usually broadly 
fused with reniform spot; reniform spot large, it and orbicular spot yellow gray, partially to 
almost completely filled with and outlined by white or light yellow scales. Hindwing: 
medium to dark fuscous with red tint; with faint median shade and discal lunule. Ab- 
domen: light gray to reddish brown. Female: similar to male. 

Male genitalia (Fig. 9). Uncus cylindrical at base, broadest at apex; with hairlike se- 
tae. Anellus heavily sclerotized laterally, covered apically with short stout spines. Juxta tri- 
angular with short dorsomedian extension. Valve 6x as long as wide, constricted near mid- 
dle; apex of valve drawn to a blunt point, without corona; sacculus 1/2 as long as valve and 
extending to dorsal margin of valve; clavus absent; ampulla of clasper gently curved, ex- 
tending posteriorly without reaching dorsal margin of valve; basal 2/3 of digitus fused to 
valve, distal 1/3 free, finger-like. Vesica approximately 1.5x as long as aedeagus, with sim- 
ple and large bilobed subbasal diverticula; single subbasal cornutus, slightly longer than 
aedeagus width, gently curved; distal 2/3 of vesica U-shaped; vesica projecting ventrally at 
base, then bending through 180° arc to project dorsally. 

Female genitalia (Fig. 13). Corpus bursae ovoid, lacking signa. Appendix bursae 
arising from posterior end of corpus bursae on right, shorter than corpus bursae, broad, 
slightly U-shaped posteriorly (apparently curving back on itself) with ductus seminalis at 
posterior end. Ductus bursae broad, dorsoventrally flattened, heavily sclerotized, scleroti- 
zation of ventral wall split by membranous cleft, ventral wall of ductus bursae produced 
ventrad into a small pouch at junction with corpus bursae. Anterior apophysis 1/4 length 
of posterior apophysis. Ovipositor lobes triangular, covered with short and long setae. 

Type specimens. Holotype: 6: WASHINGTON: Grays Harbor Co.: North Bay Bog, 
0.6 mi N of the North Bay of Grays Harbor, 47.05°N 124.09°W, elev. 5 m, 12 IV 1991, leg. 
L. G. Crabo, Native cranberry bog. Paratypes: 52 6: OREGON: Clatsop Co.: Coastal 
Plain, Ocean Home Farm 1.3 mi N of Gearhart, 46.04°N 123.90°W, Elev. 25’, 24 III 1993, 


242 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


L. & A. Crabo leg., bog E of last inland dunes (1 6). WASHINGTON: Type locality: 30 III 
1991, L. G. Crabo (5 2), 12 IV 1991, L. G. Crabo (16 3), 26 III 1993, if Troubridge (tS), 
1 IV 1994, J. & L. Troubridge (15 d); Grays Harbor Co.: 0.6 mi NW of Carlisle on Ocean 
Beach Rd., 47.16°N 124.10°W, elev. 20 m, 15 IV 1990, L. G. Crabo, native cranberry bog 
(15 3). The holotype is in the Canadian National Collection (Ottawa). Paratypes are in the 
Canadian National Collection (Ottawa), University of California (Berkeley), University of 
California (Davis), Oregon State University (Corvallis), and the personal collections of 
Lars Crabo (Bellingham, Washington), Ron Robertson (Santa Rosa, California), Jon Shep- 
ard (Nelson, British Columbia), and Jim Troubridge (Langley, British Columbia). 

We restrict the type series to specimens from Washington and Oregon because of slight 
differences in superficial appearance and habitat preferences between these and the Cali- 
fornia populations. Most California specimens are slightly smaller (forewing length: 14-15 
mm) and have more dark filling of the forewing spots than the northern populations. Also, 
some specimens from San Francisco Bay and Inverness lack most of the black markings 
and have a dark orange-brown ground color. 

Diagnosis: This is the most distinctive species in the C. cornuta group; it can usually 
be recognized without dissection. Males have narrow antennae, 2X as wide as the central 
shaft, while those of other species are 3x as wide. It is the largest species. Most specimens 
can be recognized by subtle features of wing maculation: prominent white filling of the 
forewing orbicular and reniform spots; dark reddish brown forewing ground color; promi- 
nent scalloping of the forewing lines; and conspicuous black wedges in the subterminal 
area of the forewing. Males can be identified by several genitalic features: juxta with a 
short dorsomedial extention (triangular in the other species); ampulla of clasper short, not 
reaching the costal margin of the valve (extending beyond the valve in the other species); 
vesica with one large bilobed and one simple basal diverticulum (two simple diverticula in 
the other species). Females have an appendix bursae which is smaller than the corpus bur- 
sae with the ductus seminalis joining at its posterior end. In other species the appendix 
bursae is larger than the corpus bursae with the ductus seminalis at its anterior end. Some 
indiviuals of C. gloriosa from California (vicinity of San Francisco Bay) are lighter in color 
with reduced black shading, and they lack white filling of the forewing spots. At this local- 
ity, C. gloriosa might be confused with C. cornuta, however, the two species can be differ- 
entiated by characters of the male antennae and genitalia described above. 

Distribution and biology. C. gloriosa occurs in two areas on the Pacific Coast: the Pa- 
cific Northwest (Clatsop County, Oregon and Grays Harbor County, Washington) and cen- 
tral and northern California (Humboldt, Mendocino, Sonoma, Marin, Napa, Santa Cruz, 
and Monterey Counties). Nearly all populations are located near the coastline. In Washing- 
ton and Oregon it occurs extremely locally in sphagnum bogs within ten miles of the Pacific 
Ocean. This species has not been found in sphagnum bogs in Pierce or Skagit Counties, 
Washington or near Vancouver, British Columbia. These bogs differ from those in which C. 
gloriosa occurs in that they are located in an area that was glaciated during the Pleistocene. 
The types of coastal bogs where C. gloriosa occurs might have been more widespread when 
the sea level was lower, possibly accounting for the current extended distribution of this 
species along the West Coast. C. gloriosa probably occurs further north on Washington’s 
Olympic Peninsula and possibly also in British Columbia. The California populations not 
restricted to bogs but occur in mesic forests (Ron Robertson, personal communication). 

C. gloriosa is sympatric with the other three species in the C. cornuta group, but is the 
most localized and least common. Adults fly in January to April in California and in March 
and April in the Pacific Northwest. Both sexes are attracted to lights. The early stages are 
unknown. The species was recently discovered in California by Ron Robertson and in 
Washington by the senior author. Additional California specimens were found among 
specimens of C. cornuta in museum collections. 


Cerastis enigmatica Lafontaine & Crabo, new species 
(Figs. 3, 4, 10, 14) 


Description. Male (Fig. 3): Head: reddish brown to dark gray brown; palpi reddish 
brown to dark gray brown; third segment lighter. Eyes: black. Antennae: bipectinate, 3x as 


VOLUME 51, NUMBER 3 243 


Fics. 13-16. Female genitalia of Cerastis spp. 13, C. gloriosa Crabo & Laf., Mill Val- 
ley, California, CNC 10458; 14, C. enigmatica Laf. & Crabo, Duncan, British Columbia, 
CNC 110445; 15, C. cornuta (Grt.), Mill Valley, California, CNC 10446; 16, C. robertsoni 
Laf. & Crabo, Gaviota Pass, Santa Barbara County, California, CNC 11137. 


244 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


wide as central shaft. Thorax: reddish brown to gray brown; collar lighter, buff to reddish 
tan, with darker transverse lines. Foretibiae: spined. Forewing: length: 13-15 mm, ground 
color variable, reddish tan to gray brown, with lighter tints, suffused with variable amounts 
of black in cell; basal area frequently paler or with more extensive gray scaling than medial 
or subterminal areas; terminal area paler reddish brown than remainder of forewing; lines 
double, black with pale filling; basal line faint, sinuous; antemedial line oblique, gently ex- 
curved, notched basad in fold; median shade usually obscure, when evident a wavy line ex- 
tending from reniform spot to inner margin; postmedial line scalloped, excurved opposite 
reniform spot, lower half nearly straight; subterminal line weak, pale, undulating, with 
slender black wedges proximally near costa on radial veins and opposite cell between me- 
dian veins; terminal line thin, weakly scalloped; claviform spot oblong, black; orbicular 
spot large, gray or brown, an elongate ellipse, broadly fused with reniform spot; reniform 
spot large, gray or brown, it and orbicular spot outlined with yellow scales. Hindwing: 
medium to dark fuscous, with darker median shade and discal lunule. Abdomen: light gray 
to reddish brown; untufted, but with long hairlike scales on tergum one. Female (Fig. 4): 
similar to male. 

Male genitalia (Fig. 10). Similar to those of C. cornuta, except for shape of vesica. 
Vesica approximately 1.5x as long as aedeagus; distal 2/3 strongly curved, extending first 
towards left. Two simple subbasal diverticula present. Single cornutus straight. 

Female genitalia (Fig. 14). Similar to C. cornuta except for bursa shape. Appendix 
bursae joined to ventral corpus bursae 1/3 from posterior end, extending toward corpus 
bursae anteriorly. 

Type specimens. Holotype: 3: British Columbia: Vancouver, 22 III 1903, ex. coll. 
Bush Wilson. Paratypes: 145 5, 19 9: ALASKA: Ketchikan, 24—29 IV (1 d). BRITISH CO- 
LUMBIA: Vancouver Island: Wellington, 2 IV 1903, 27 III 1906 & 4—14 II 1906, Coll. G. 
W. Taylor (935,52), 251V 1903 &8 & 16 IV 1904, Bryant (5 5, 1 2), 29 III 1949 & 10 IV 
1949, Woodcock (3 3); Vancouver Island: Quamichan, 27—28 III 1907, 28 III 1908 & 4 IV 
1909, G. B. Day (2 4, 2 2); Vancouver Island: Duncans, 27 III 1908, 8 IV 1908 & 1-7 IV 
1908, Hanham (1 d, 2 2); Vancouver Island: Mill Bay, 14 VI 1986, K. B. Bolte (1 d); Van- 
couver, 23 III 1903, ex. coll. Bush Wilson (1 3), 3 IV 1904 (1 3), 192_, W. Downes (2 4); 
Langley, 5 km E, 1-7 IV 1991 & 1-4 IV 1992, J. Troubridge (5 4); Queen Charlotte Is- 
lands: Massett, 28 V 1894, J. H. Keen (1 4); Skagit River Valley, near N end of Ross Lake, 
9 IV 1994, J. Troubridge (14 ¢); 16 IV 1994, J. Troubridge and A. & L. Crabo (15 4, 2 9). 
WASHINGTON: Cowlitz Co.: Columbia R. valley, 3 mi N of Kalama, SW Carroll's Bluff, 
46.05°N 122.86°W, 50 m, 13 IV 1991, L. & A. Crabo leg., disturbed hillside with oak (1 3); 
Cowlitz R. valley, Toutle Rest Area on Interstate 5, 46.35°N 122.90°W, 360’, 24 III 1993, 
L. & A. Crabo leg., lowland forest (1 5); Grays Harbor Co.: Ocean City, 26 HI 1993, J. 
Troubridge (1 3); North Bay Bog, 0.6 mi N of North Bay of Grays Harbor, 47.05°N 
124.09°W, elev. 5 m, 12 IV 1991, L. G. Crabo leg., Native cranberry bog (1 4), 26 III 1993, 
J. Troubridge (5 4), 1 IV 1994, J. & L. Troubridge (3 4); Humptulips R. valley, Copalis 
Crossing, 47.10°N 124.07°W, 20 m., 30 III 1992, L. G. Crabo leg., Storefront lights (1 d); 
[Island Co.]: Deception Pass, 27 III 1993, J. Troubridge (1 3), 2 IV 1994, J. & L. 
Troubridge (1 ¢); King Co.: Factoria, 9 IV 1949, E. C. Johnson (1 4, 1 °); 7.5 mi E of North 
Bend on Middle Fork Snoqualmie R., 300 m, 47.50°N 121.63°W, i3 IV 1988, leg. L. 
Crabo (2 4, 2 2); Kitsap Co.: Bremerton, 1 IV 1948, Don Frechin (1 2); Klickitat Co., Co- 
lumbia River Gorge, Major Creek 1/2 mi N of Columbia River, 9 IV 1994, L. G. Crabo (1 
3); Mason Co.: Shelton, 16 IV 1949, E. C. Johnson (4 2); 2 mi E Little Hoquiam, Grape- 
view Loop Rd., 25 m, 47.31°N 122.90°W, 24 III 1990, L. Crabo leg. (4 3): Elfendahl Pass 
Rd. 0.3 mi N of Hwy. 302, 2 mi W of Belfair State Park, 47.42°N 122.91°W, elev. 50 m, 29 
III 1989, L. Crabo leg. (11 3); Okanogan Co., Early Winters canyon, Highway 20 1 mi S$ of 
Lone Fir Campground, 7 V 1994, L. & A. Crabo and C. Coughlin (8 3); Pierce Co.: Puget 
Trough, NE corner of Cranberry Lake, 46.90°N 122.36°W, 644’, 11 III 1992, L. Crabo 
leg., native sphagnum bog (6 2); [San Juan Co.]: Orcas Island, 18 IV 1949, E. Hendriksen 
Coll. (1 3); Skagit Co., 3.5 mi SE of Big L., Cavanaugh Rd. at Grandstrom Rd., 48.32°N 
122.16°W, 550’, 14 IV 1993 L. G. Crabo, sphagnum bog (1 4); Snohomish Co.: S. Lake 
Ballinger, 47.95°N 122.32°W, 25 m, 3 III 1988, leg. L. Crabo (2 3); Thurston Co.: 3 mi N 
Tenino, Rocky Prairie, 50 m., 46.89°N 122.87°W, 25 III 1990, leg. L. Crabo (2 3); What- 


VOLUME 51, NUMBER 3 QA5 


com Co.: Chuckanut Bay of Bellingham Bay, elev. 35 m, 48.69°N 122.49°W, 19 IV 1993, 
L. G. Crabo leg., dry rock slope (1 ¢); Skagit River Valley near N end Ross Lake, 16 IV 
1994, J. Troubridge and A. & L. Crabo (4 3); Mt. Baker Hwy. at N Fork Nooksak River 
crossing, 1/2 mi N of Silver Fir Campground, 23 IV 1994, L. Crabo (19 d, 4 2); Yakima Co., 
Tieton River Valley, Oak Creek at Tieton R., 46.72°N 120.81°W, 550 m, 18 IV 1992, L. G. 
Crabo, riparian forest with oak (1 3). 

The holotype is in the Canadian National Collection (Ottawa). Paratypes are in the 
Canadian National Collection (Ottawa), University of California (Berkeley), University of 
California (Davis), Oregon State University (Corvallis), and the personal collections of 
Lars Crabo (Bellingham, Washington), Ron Robertson (Santa Rosa, California), and Jim 
Troubridge (Langley, British Columbia). 

We restrict the type series to specimens from Washington, British Columbia, and 
Alaska. 

Diagnosis. Cerastis enigmatica is most similar to C. cornuta and C. gloriosa. It is 
sympatric with C. gloriosa, but can be separated from it by features described in its diag- 
nosis. The male genitalia of C. enigmatica differ from those of C. cornuta and C. gloriosa 
by the shape of the vesica, which is strongly curved distally, not straight. Females differ 
from these two species in that the appendix bursae joins the corpus bursae 1/3 from its 
posterior end, not at the posterior end. Adults are nearly identical to C. cornuta, which oc- 
curs to the south of the range of C. enigmatica. Cerastis enigmatica adults tend to be 
slightly larger with broader wings, they have more contrasting forewing maculation, more 
prominent light borders surrounding the forewing spots, and slightly lighter hindwings. 

Distribution and biology. Cerastis enigmatica is moderately common to abundant 
in mesic conifer forests at low elevations in the Cascade Mountains and on the west coast 
from the Alaska Panhandle to southern Humboldt County, California (Miranda). It also 
occurs on the east slope of the Cascades but is much less common there. It occurs with C. 
gloriosa in Washington, Oregon, and at Willow Creek, Humboldt County, California. The 
adults are active from March to late April at the time when most deciduous trees are in 
bloom. Both sexes are attracted to light. The early stages are unknown. 

C. enigmatica is moderately common in collections but has until now been confused 
with C. cornuta. All previous records of C. cornuta from Oregon, Washington, Alaska, and 
British Columbia are C. enigmatica. 


Cerastis cornuta (Grote) 
(Figs. 5, 6, 11, 15) 


Pachnobia cornuta Grote, 1874, Bull. Buffalo Soc. Nat. Sci., 2:68. 
Type Locality: |California, USA]. [Lectotype in BMNH] 


Type specimens. Pachnobia cornuta was described from “two fresh specimens” 
stated to be in the Collection of the Buffalo Society of Natural Sciences. A number of spe- 
cies said to be in that collection (e.g., Agrotis specialis Grote, Agrotis formalis Grote) have 
not been found in the collection of the Buffalo Museum of Science but specimens from 
the Grote Collection labeled “type” and exactly matching the original description are in 
the Natural History Museum, London. For other species (e.g., Agrotis wilsoni Grote) the 
nominal “type” in The Natural History Museum differ significantly from the description 
and the types must be considered to be lost. For Pachnobia cornuta Grote, a male in The 
Natural History Museum, London, purchased from the Grote Collection and labeled as a 
type, matches the description of the species in every detail and is hereby selected as lec- 
totype. The specimen is labeled “Type/ California Grote Coll. 81-116/ Metalepsis cornuta 
Grote/ Metalepsis cornuta Grote Type.” 

Diagnosis. Adults of C. cornuta (Fig. 5, 6) are nearly indistinguishable from C. enig- 
matica and C. robertsoni and are similar to some specimens of C. gloriosa. The male gen- 
italia of C. cornuta (Fig. 11), C. enigmatica, and C. robertsoni are indistinguishable except 
for the shape of the vesica. All three differ from C. gloriosa by the presence of a triangular 
juxta without median extension, longer claspers cine extend beyond the dorsal margin of 
the valves, and the presence of two simple diverticula on the vesica. The distal portion of 


246 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the vesica of C. cornuta differs from those of C. robertsoni and C. enigmatica in being 
straight beyond the basal turn. Females of these three species are distinguished from C. 
gloriosa by having an appendix bursae which is larger than the corpus bursae, with the 
ductus seminalis joining it at its anterior end. The female genitalia of C. cornuta (Fig. 15) 
differ from those of C. enigmatica in that the appendix bursae joins the corpus bursae at 
its posterior end. They differ from those of C. robertsoni in that the corpus bursae is con- 
stricted posteriorly, the appendix bursae is even in width throughout its length and pro- 
jects obliquely away from the corpus bursae anteriorly, and the ductus bursae is shorter, 
1.5x as long as wide. Cerastis cornuta differs from C. enigmatica in slightly smaller size 
(forewing length: 11-13 mm versus 13-15 mm), less contrasting forewing pattern, and less 
prominent yellow outline of the orbicular and reniform spots. Cerastis robertsoni is 
slightly smaller (forewing lenth: 10-13 mm) but is difficult to separate from C. cornuta 
without dissection. 

Distribution and biology. Cerastis cornuta is moderately common in forests in 
western California from Sonoma County south to Santa Clara County. It, like C. gloriosa, 
occurs near the Pacific Ocean with most records from the vicinity of San Francisco Bay. It 
occurs with C. gloriosa at a few locations. The adults are active from mid-January through 
late April. The early stages are unknown. 


Cerastis robertsoni Lafontaine and Crabo, new species 
(Figs. 7, 8, 12, 16) 


Description. Male (Fig. 7): Head and palpi: appearing chestnut (actually a mixture 
of white, buff, brown, and black scales). Eyes: black. Antennae: bipectinate, 3x as wide as 
central shaft. Thorax: dark brownish gray; collar pale brownish gray, contrasting with 
darker head and thorax. Foretibiae: spined. Forewing: length: 10-13 mm; ground color 
light to dark brownish gray, suffused with variable amounts of black in discal cell; basal and 
terminal areas and costa paler gray; lines double, black, pale-filled; basal line sinuous; an- 
temedial line oblique, gently excurved, notched towards wing base in fold; median shade 
wavy, evident only from reniform spot to inner margin of forewing; postmedial line scal- 
loped, lower half nearly straight, excurved opposite discal cell; subterminal line weak, pale, 
undulating, with slender black wedges proximally between veins M1 and M2 and near 
costa on radial veins; terminal line thin, weakly scalloped; claviform spot narrow, black; or- 
bicular spot large, variable, gray, brown, or white, variably outlined with thin pale line, 
broadly fused with reniform spot and often extended onto forewing costa; reniform spot 
narrow, oblique, angled towards orbicular spot, gray with partial black and paler gray out- 
line. Hindwing: dark fuscous, with darker discal lunule and slight trace of median line. Ab- 
domen: gray with long brownish gray hairs; untufted but with numerous long hairs on an- 
terior margin of tergum one. Female (Fig. 8): similar to male. 

Male genitalia (Fig. 12). Similar to C. cornuta except for size of aedeagus and shape 
of vesica. Aedeagus smaller than that of C. cornuta, but with same proportions: vesica ap- 
proximately 1.5x as long as aedeagus. Vesica bending sharply ventrad 90°, then bent 90° to 
project to left, distal 2/3 nearly straight. Two simple subbasal diverticula present, one large 
and one small. Single cornutus straight. 

Female genitalia (Fig. 16). Similar to C. cornuta except for shape of corpus bursae 
and length of ductus bursae. Corpus bursae broadly fused with appendix bursae, without 
posterior constriction. Appendix bursae constricted apically; apical portion curving dor- 
sally to project dorsad. Ductus bursae 2x as long as wide. 

Type specimens. Holotype: 6: CALIFORNIA: Monterey Co.: Big Creek Res., UC- 
NLWBRS, 2 II 1994, L. Crabo, J. Powell, & R. Robertson. Paratypes: 38 3, 10 2: CALI- 
FORNIA: Monterey Co.: Chualar, 11 I 1963 (1 3); Big Creek Res., UCNLWRS, Trail to 
Redwood Camp, 80 m, 24/26 I 1988, J. A. Powell (5 3); Big Creek Res., UCNLWRS, So. 
Ridge Rd., 220 m, bl. trap, 24 I 1988, J. A. Powell (1 4), 28 II 1989, J. Powell & M. Pren- 
tice (1 d, 1 °); Big Creek Res., UCNLWBRS, So. Ridge Pt., 275 m, bl. trap, 24/26 I 1988, J. 
A. Powell (1 3); Big Creek Res., UCNLWRS, So. Gate Rd., 190-200 m, bl., 24/26 I 1988, 
J. A. Powell (4 d), 21/22 II 1988, J. A. Powell (1 ¢), 28 II 1989, J. Powell & M. Prentice (al 
3); Big Creek Res., UCNLWRS, HQ area, 1-10 m, coastal scrub, bl. trap, 24/26 I 1988, J. 


VOLUME 51, NUMBER 3 247 


A. Powell (3 d, 2 2), 28 II 1989, J. Powell & M. Prentice (1 2); Big Creek Res., UCNL- 
WRS, 2 II 1994 L. G. Crabo, J. Powell, & R. Robertson (15 d, 4 2); Santa Barbara Co.:: 
Gaviota Pass, 3 February 1987, Powell & Wagner, blacklight (5 d, 2 2). An additional 6 
males and | female from the type locality and Gaviota Pass were examined but were ex- 
cluded from the type series because of their poor condition. 

The holotype is in the University of California (Berkeley). Paratypes are in the Cana- 
dian National Collection (Ottawa), the University of California (Berkeley), the University 
of California (Davis), and the personal collections of Lars Crabo (Bellingham, Washing- 
ton) and Ron Robertson (Santa Rosa, California). 

Diagnosis: Cerastis robertsoni is the smallest species of Cerastis in North America. 
It is most similar to C. cornuta, which occurs in central California to the north of the range 
of C. robertsoni. Cerastis robertsoni tends to have more gray in the forewing, and less con- 
spicuous light-colored outlines of the forewing spots. However, there are no reliable char- 
acters for separating these species without dissection. Males of C. robertsoni differ from 
those of C. cornuta and C. enigmatica by the shape of the vesica: the distal portion of the 
vesica in C. robertsoni extends straight toward the left beyond two 90° basal bends. The 
female genitalia are most similar to those of C. cornuta, but differ by several features: the 
corpus bursae is broad without posterior constriction; the appendix bursae is broadly fused 
with the corpus bursae posteriorly and is constricted apically, curving dorsally; and the 
ductus bursae is longer (2x as long as wide). They differ from those of C. enigmatica by 
having the corpus bursae and appendix bursae joined at their posterior ends. Cerastis 
robertsoni is unlikely to be confused with C. gloriosa. However, some specimens have the 
forewing spots filled with pale yellow, and might mistakenly be identified as C. gloriosa us- 
ing our key. 

Distribution and biology. Cerastis robertsoni is known only from three localities 
but may occur more broadly in the Coast Range of central and southern California from 
Monterey Bay to Santa Barbara County. It is moderately common in California coastal 
chapparal habitat (J. Powell, pers. comm.) but has seldom been collected, probably be- 
cause of its flight period early in the year. It is sympatric with C. gloriosa at the type local- 
ity in Monterey County. Adults are active in January and February. Both sexes are attracted 
to light. The early stages are unknown. 

All but one of the known specimens of C. robertsoni were collected by Jerry Powell and 
his field associates. It was recognized as distinct from C. cornuta by Ron Robertson who 
sent specimens to the junior author for evaluation. We take pleasure in naming this spe- 
cies after Mr. Robertson in recognition of his contribution. 


ACKNOWLEDGMENTS 


Paul C. Hammond (Oregon State University, Corvallis), Steven Hayden (University of 
California, Davis), Martin R. Honey (The Natural History Museum, London), Robert W. 
Poole (National Museum of Natural History, Washington D.C.), Jerry A. Powell (University 
of California, Berkeley), Frederick H. Rindge (American Museum of Natural History, New 
York), Eric H. Metzler (Columbus, Ohio), Ronald Robertson (Santa Rosa, California), and 
James T. Troubridge (Langley, British Columbia) made specimens in their care available 
for study. Michael Fibiger provided information on the genitalia of Eurasian species of 
Cerastis. Eugenie Krelina prepared the genitalic dissections and William Lukey photo- 
graphed the adults and genitalia. Jonathan P. Pelham, Jerry A. Powell, and Ronald Robert- 
son accompanied the senior author to collect parts of the type series of C. gloriosa and C. 
robertsoni. Neale Maine (Gearhart, Oregon) gave permission to collect C. gloriosa on his 
property and graciously provided lodging for the freezing collectors. Jonathan P. Pelham, 
Rodney Crawford, and James T. Troubridge critiqued early versions of the manuscript. 


LITERATURE CITED 
FIBIGER, M. 1993. Noctuidae Europaeae, Vol 2, Noctuinae II. Entomological Press. Soro. 
Denmark. 230 pp. 
FRANCLEMONT, J. G. & E. L. TODD. 1983. Noctuidae. In Hodges, R. W. et al. (Eds.), 
Check list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd. and The 
Wedge Entomol. Research Foundation. London. 284 pp. 


248 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


HOopDGEs, R. W. 1971. Sphingoidea. In Dominick, R. B., et al., Moths of America north of 

Mexico. Fascicle 21. E. W. Classey Ltd. and R. B. D. Publications Inc. xii + 216 pp. 14 
Is. 

Moe eR ee ee J. 1927. Notes on certain Agrotid genera and species (Lepid.). Canadian 
Entomol. 59:64—66. 

. 1938. Check list of the Lepidoptera of Canada and the United States of America. 
Part 1. Macrolepidoptera. Mem. Southern Calif. Acad. Sci. 1:1-272. 

POOLE, R. W. 1989. Lepidopterorum Catalogus (New Series). Fasc. 118. Noctuidae. Lei- 
den. E. J. Brill. 1314 pp. 


Received for publication 3 March 1996; revised and accepted 17 July 1996. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
51(3), 1997, 249-256 


NOTES ON THE SESIID FAUNA OF SOUTHWESTERN WEST VIRGINIA 
Additional key words: pheromones, faunal surveys, Appalachian region. 


The Sesiidae is a relatively well studied moth family. In the past century, three compre- 
hensive monographs have been written on the group (Beutenmuller 1901, Engelhardt 
1946, Eichlin & Duckworth 1988), and several regional surveys have been conducted in 
the eastern United States (Neal et al. 1983 in Maryland, Sharp et al. 1978 in Florida, 
Sharp et el. 1979 again in Florida, Snow et al. 1985 in Georgia, Solomon et al. 1982 in Mis- 
sissippi, and Taft & Snow in 1991 in the north central United States). These studies were 
aided enormously by the chemical isolation and synthesis of female sesiid sex attractants. 
In 1974 the synthesis of the (Z,Z)-3,13-octadecadien-1-ol acetate and of the (E,Z)-3,13-oc- 
tadecadien-1-ol acetate was achieved by Tumlinson and Yonce respectively, and in 1983 the 
(E,Z)-2,13-octadecadien-1-ol acetate was isolated by Schwartz (Eichlin & Duckworth 1988). 
The use of these pheromones, alone or in different combinations, in the acetate or alcohol 
forms, has become the principal method for collecting male Sesiidae and has led to the dis- 
covery of many new species and to the elucidation of distributions and flight patterns. 

The purpose of the present study is to establish the number of Sesiidae species flying in 
southwestern West Virginia, and to report on male responsiveness to different synthetic 
pheromone isomers in “pure” form and in different combinations. 

Sampling was conducted over a period of five years, starting in May 1990 and ending in 
September 1994. Seven locations in two counties were sampled: five in Kanawha County 
and two in adjacent Putnam County. Three of the Kanawha County locations are situated 
in the city of Charleston (South Hills, Kanawha City, Coonskin Park). Kanawha State For- 
est is located at the southern boundary of the city, and Tupper Creek is a small rural com- 
munity 10 miles north of the city. The South Hills and the Kanawha City locations are 
densly populated urban areas, with occasional patches of oak woodland. Coonskin Park is 
a partially developed hilltop that is well drained and rather dry. In addition to the decidu- 
ous trees typical for the area, it contains a large population of old pines. Kanawha State 
Forest is a large tract of undeveloped land containing a variety of microhabitats including 
gulches, ridges, river bottoms and hills with large tracts of deciduous forest interrupted by 
small clearings. Tupper Creek is a disturbed area along a small creek with an extensive 
growth of willow trees. Of the two Putnam County locations, one is situated 20 miles west 
of Charleston on a fir tree farm, and is surrounded by hills of uninterrupted deciduous for- 
est. The other site, located 25 miles west of Charleston, is a disturbed area on the south 
bank of the Kanawha River, which supports an extensive growth of willow. 

I used Multipher I traps and lures supplied by the IPM Great Lakes Company 
(Vestaburg, MI). The lures used in this study are listed in Table 1. In the traps I placed 
DDVP toxicant strips provided by IPM Great Lakes. In 1990, five traps baited with 97:3, 
ZZA, EZA, 1:1, and TRI lures were deployed in the South Hills area of Charleston. In 
1991, eight traps baited with 97:3, ZZA, EZA, TRI, EZ-2,13-OH, EZ-2,13-A, EZ-3,13- 
OH, and 99:1 were used on the fir farm in Putnam County. In 1992, 2. eight traps baited 
with 97:3, EZA, ZZA, 1:1, TRI, 99:1, and EZ-3,13-OH were used in Kanawha State For- 
est. In 1993, five traps baited with 97:3, ZZA, EZA, 1:1, and 99:1 were used at Tupper 
Creek and three traps with 97:3, EZA, and ZZA in Kanawha State Forest. In mid June 
1993, the 1:1 and 97:3 traps were moved from Tupper Creek to South Hills and the 99:1 
trap to Kanawha State Forest. Then, in mid August all traps were moved to Kanawha City. 
In 1994, three traps baited with 97:3, ZZA, and EZA were used on the Kanawha River 
shore; three traps baited with 97:3, ZZA, and EZA in Coonskin Park and two traps baited 
with 1:1 and 2:1 in Kanawha City. At the beginning of August, the three Putnam County 
traps were moved to Kanawha City. 

I began each baiting season in May, when the weather was reliably warm, and ended 
trapping in late September or early October. In the middle of July I added a new lure load 
to each trap, making sure that each trap carried the same lure throughout the season; new 


250 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Pheromonal lures used in the present study and their abbreviations. Small 
percentages of impurities are omitted (these should be less than 1% with current synthe- 
sis techniques; Tumlinson 1979). 


—————————— 
Abbreviation Pheromone 


97:3 97% (Z,Z)-3,13-octadecadien-1-ol 
acetate/3%(E,Z)-3, 13-octadecadien-1-ol acetate 

ZZA 100% (Z,Z)-3,13-octadecadien-1-ol acetate 

EZA 100% (E,Z)-3,13-octadecadien-1-ol acetate 

lel 50:50 mixture of (Z,Z)-3,13-octadecadien-1-ol acetate/ 
(E,Z)-3,13-octadecadien-1-ol acetate 

99:1 99:1 mixture of (E,Z)-2,13-octadecadien-1-ol acetate/ 
(Z,Z)-3, 13-octadecadien-1-ol acetate 

EZ-2,13-OH 100% (E,Z)-2,13-octadecadien-1-ol 

EZ-2,13-A 100% (E,Z)-2,13-octadecadien-1-ol acetate 

EZOH 100% (E,Z)-3,13-octadecadien-1-ol 

TRI 20:1:3 mixture of (Z,Z)-3,13-octadecadien-1-ol acetate/ 


(E,Z)-3, 13-octadecadien-1-ol/acetate 
(Z,Z)-3, 13-octadecadien-1-ol 
9:1 2:1 mixture of (E,Z)-3,13-octadecadien-1-ol acetate/ 
(E,Z)-3,13-octadecadien-1-ol 


toxicant vapor tapes were added at the same time. Traps were placed on tree branches, ei- 
ther at the edge of the forest or on solitary trees, in clearings, at 1-1.5 m above ground 
with at least 20 m distance between traps. Whenever possible the traps were placed in a 
south or southwest orientation. I visited each trap once a week and emptied the buckets 
into labeled styrofoam cups that I took home to record the attracted specimens. 

During this study 5080 male sesiids were captured, representing 24 species in 8 genera. 
Two of these, Synanthedon kathyae Duckworth & Eichlin and Alcathoe carolinensis En- 
gelhardt, are new records for the state of West Virginia (T. D. Eichlin, pers. comm.). 
Melittia cucurbitae (Harris) was never lured to a pheromone trap but was netted on the 
foodplant (zucchini squash); since the main purpose of this study was to record as many 
species as possible for this area, I included it in the present listing. Table 2 lists all the 
recorded species, showing their yearly abundance at the sample sites. Table 3 gives the 
monthly distributions and abundances, and Tables 4—5 summarize the sensitivity of male 
sesiids to the different pheromone lures in this study. 

Geographical and temporal trends. The records of Synanthedon pictipes (Grote 
& Robinson), Podosesia aureocincta Purrington & Nielson, Carmenta bassiformis 
(Walker) and Synanthedon rileyana (Hy. Edwards) show wide year to year fluctuations. 
Since these four species have at least one generation per year (S. pictipes is multivoltine), 
these fluctuations were probably geographical rather than temporal: one year the traps 
happened to be placed inside the territory of a population and attracted a large number of 
individuals; another year they were outside the population’s territory and attracted fewer 
individuals. The number of attracted specimens suggests that these species are abundant 
inside well circumscribed colonies, which are doubtless centered around their foodplants. 

Synanthedon rubrofascia (Hy. Edwards), Synathedon scitula (Harris), Synanthedon de- 
cipiens (Hy. Edwards), Synanthedon fatifera Hodges and Carmenta ithacae (Beuten- 
muller) had similar yearly/geographical fluctuations, but their numbers suggest much 
smaller and even more circumscribed colonies. It is worth noting the difference in abun- 
dance between Synanthedon exitiosa (Say) and S. pictipes. While these species both uti- 
lize wild and cultivated Rosaceae as hosts (Snow 1985, Eichlin 1988), S. pictipes appears 
to be much less abundant and more localized in southwestern West Virginia than S. exi- 
tiosda. 

The records for Paranthrene simulans (Grote) and Paranthrene pellucida Greenfield & 
Karandinos confirm their two-year life cycle, with peak numbers being attracted in odd 
years. In the study area, both species appear to have similar abundance, both peak in the 


VOLUME 51, NUMBER 3 951 


TABLE 2. Yearly occurence of sesiid species in southwestern West Virginia based on 
captures in pheromone baited traps. 


Species Total 1990 1991 1992 1993 1994 
Podosesia syringae 1564 67 613 176 488 220 
Synanthedon exitiosa 1229 alte 168 159 353 336 
Podosesia aureocincta 651 3 93 547 i Il 
Synanthedon pictipes 519 113 142 9) 4] 214 
Carmenta bassiformis 239 6 2 56 147 24 
Paranthrene simulans 181 0) 89 4 87 il 
Paranthrene pellucida 170 5 95 2 65 3 
Alcathoe caudata Dz 120 0) 5 2 0 
Synanthedon rubrofascia 115 33 3 1 Ia 61 
Synanthedon rileyana 110 2) 105 2 i O 
Synanthedon scitula Be} 1 25 25 il 1 
Synanthedon fatifera 29 2 14 0 0 1s} 
Carmenta ithacae 20 @) 3 O 0) iY 
Synanthedon acerni 13 O 2 2 O 9 
Synanthedon acerrubi 1; 0) itil IL @) 0) 
Synanthedon decipiens 1) O 0 0) D 10 
Sannina uroceriformis 12 0 12 0 0) 0) 
Synanthedon viburni Il 3 0 1 0 7 
Synanthedon rhododendri 8 0 0) il 0 1 
Synanthedon kathyae 3 0) 0) 3 0) 0) 
Vitacea polistiformis 2 0) 0 2 0) 0) 
Alcathoe carolinensis 2 O O 1 0) Il 
Vitacea scepsiformis 1 0) 0 0 1 0) 
Mellittia cucurbitae 1 0 0) 0 it 0 


same years, and their two-year cycles occurred in odd numbered years, contrary to Engel- 
hard’s statement (Engelhardt 1946:146) that even numbered years have peak flights of P. 
simulans in eastern United States. 

Table 3 shows that the most productive months were June and July. All but one of the 24 
species were caught during these two months, the exception being the later-flying P. aure- 
ocincta. Table 3 also shows distinct temporal segregation of Podosesia syringae (Harris) 
and P. aureocincta, the first having peak flight in May and the latter reaching peak flight in 
September. This lack of overlap is similar to other reports (Eichlin & Duckworth 1988) of 
an April-May peak for P. syringae and a peak after July for P. auroecincta. The closely re- 
lated P. simulans and P. pellucida also showed a segregation in flight periods. P. simulans 
had peak flight in May, wheras P. pelludica peaked in June. Unfortunately, due to unfavor- 
able weather, I never set traps up prior to May; therefore I lack data about the responsive- 
ness of P. simulans to sex attractants in even earlier months. 

The phenology for S. pictipes suggests two generations per year, with a more abundant 
spring generation in May and June (probably starting earlier) and a less abundant summer 
generation from July through October. This species is the only one that came to the traps 
in each month of the study. S. exitiosa had an extended flight season as witnessed by other 
authors (Eichlin & Duckworth 1988, Snow et al. 1985). S. rubrofascia exhibited a similar 
pattern, with a flight period extending from May to September and a peak in July. This 
matches the data reported by Snow et al. (1985), who were able to capture the moth from 
April to November in central Georgia. Eichlin and Duckworth (1988) give a shorter flight 
period (May and June). The specimens captured in this study did not resemble the illus- 
tration given by Eichlin and Duckworth in their monograph of the Sesiidae, but, having 
completely transparent forewings, they matched the illustration given by Taft and Snow 
(1991). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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254 


TABLE 5. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


in three different studies. 


Present Eichlin & Taft & 
Species Study Duckworth, (1988) Snow, (1991) 

P. syringae 97:3, ZZA ZZA ZZA 

S. exitiosa O7-SSZZN ZZA ZZA 

P. aureocincta OVAS8), IIRL, ZAAN TRI ZZA, 50/50 ZZA/EZOH 

S. pictipes EZA EZA EZA 

C. bassiformis EZA ZZA, ZZOH ZZA, 50/50 ZZA/ZZOH 

P. simulans Os8), E74, ISR 96:4, ZZA ZZA, 99:1 

P. pellucida 97:3, ZZA no mention ZZA 

A. caudata TRI no mention ZZOH, 1:1 

S. rubrofascia 1:1, 2:1 EZA/EZOH JLeJE Tea 

‘Ss! rileyana EZ-2, 13-OH EZOH, EZOH, 50/50 
EZOH/EZA ZZA/EZOH 

S. scitula EZA ZZA ZZA 

S. fatifera ZZLA, 97:3 ZZA ZZA 

C. ithacae EZA ZZA, 97:3 EZA 

S. acerni TL biN, SUS}. light ZZA, light ZZA, 50/50 ZZA/ZZOH 

S. acerrubi 99:1 IDVZ=O. IBGE 99:1 

S. decipiens 97:3, ZZA ZZA/EZA, ZZA, 1:1 
ZZA/EZOH 

S. uroceriformis EZ-2, 13-OH EZOH, no mention 
EZOH/ZZOH 

S. viburni EZA EZA EZA 

S. rhododendri 97:3, ZZA ZZA no mention 

S. kathyae EZOH, ZZA ZZA no mention 

V. polistiformis oor 99:1 Seal 

A. carolinensis EZA, 1:1 HAAS oa no mention 

V. scepsiformis 99:1 ZZA/EZA, no mention 
ZZA/EZOH 

M. cucurbitae on foodplant 99:1 99:1 


Comparison of responsiveness of sesiid species to different pheromone lures 


Synanthedon acerni (Clemens) and Synanthedon acerrubi Engelhardt had a shorter 
flight period than given by other sources (Eichlin & Duckworth 1988, Snow et al. 1985, 
Taft & Snow 1991). Both species were active only in May and June. Sannina uroceriformis 
Walker was active in May and June (see Eichlin & Duckwoth 1988, Snow et al. 1985). 
Synanthedon viburni Engelhardt had a more extended flight period (May through August) 
than S. fatifera, which was active almost exclusively in June. S. scitula, S. rhododendri 
(Beutenmuller), S. rileyana, S. decipiens, C. bassiformis, C. ithacae and Alcathoe caudata 
(Harris) had flight periods similar to those observed by other authors (Eichlin & Duck- 
worth 1988, Snow et al. 1985, Taft & Snow 1991). A. carolinenesis had an earlier activity 
period (June and July) compared to other sources (Eichlin & Duckwoth 1988, Snow et al. 
1985, Sharp et al. 1978). This is somewhat surprising, considering that both Snow’s and 
Sharp’s groups collected their specimens to the south of West Virginia (Georgia and 
Florida, respectively). The flight of S. kathyae corresponded with the period given by 
Eichlin and Duckworth (1988), as did that of Vitacea polistiformis (Harris) and Vitacea 
scepsiformis (Hy. Edwards). 

Pheromonal responses. The 97:3 blend proved to be the most generalized attrac- 
tant, yielding a total of 15 species. It was the main lure for seven species: S. exitiosa, P. 
pellucida, P. syringae, S. decipiens, S. rhododendri, P. aureocincta and P. simulans. It also 
attracted a large percentage of all S. fatifera, S. viburni, S. acerni and S. scitula. The 
“pure” ZZA lure also attracted 15 species but was the main attractant for only two species: 
S. fatifera and S. acerni (note that 38% of S. acerni individuals were caught at black light, 
usually early in the morning; this was the only species that came to black light during this 


VOLUME 51, NUMBER 3 259 


study, see Eichlin and Duckworth 1988). The “pure” EZA pheromone was attractive to 11 
species; for six of those it was the main attractant: C. ithacae, S. pictipes, S. scitula, C. 
bassiformis, S. viburni and A. carolinensis. Surprisingly, the TRI mixture was only moder- 
ately attractive for P. aureocincta, for which it was originally formulated (Nielsen et al. 
1979), attracting only 21% of the 651 individuals of P. awreocincta caught. In contrast, 
59% of the individuals responded to the 97:3 mixture. Sharp and Eichlin report the same 
weak attraction of this combination for P. aureocincta (1979). 

The TRI lure proved strongly attractive to A. caudata (92% of the total). A. caudata 
displayed a specific pheromone responsiveness: of 120 specimens caught in 1990 in South 
Hills, 117 responded to the TRI lured traps and only three to the 97:3 traps. In 1991 no 
individuals came to the TRI trap in Putnam County. In subsequent years I did not have 
the TRI lure, but the 97:3 lure attracted 5 and 2 individuals respectively in Kanawha State 
Forest and Tupper Creek. This suggests that A. caudata occurs in large, well circum- 
scribed colonies, and the individuals exhibit a strong attraction to the TRI mixture. Synan- 
thedon acerrubi Engelhardt and Sannina uroceriformis Walker exhibited the same geo- 
graphical confinement, but the numbers caught indicate small colonies or weak responses. 
The 1:1 mixture attracted S. rubrofascia (76% of the total). The 99:1 blend lured all the V. 
polistiformis and V. scepsiformis caught and was highly attractive for S. acerrubi (83% of 
the total). EZ-2,13-OH was a highly specific lure for S. uroceriformis, (100% of the total). 
It also proved attractive to S. rileyana (83% of the total). The EZOH isomer was a less ef- 
fective attractant for S. rileyana (10% of the total), and lured two individuals of S. kathyae 
(66%). No M. cucurbitae males were attracted to any pheromone traps, even though a 
99:1 baited trap was kept close to the patch with zucchini-squash plants, where the only 
individual caught during this study was netted. 

Table 5 summarizes male sesiid responsiveness to different sex lures in three different 
studies: the present work, Eichlin and Duckworth (1988) and Taft and Snow (1991). In 
the present study C. bassiformis exhibited a strong affinity toward the EZA lure (77% of 
the total), whereas the other two studies reported the ZZA isomer as most attractive. Since 
these were the same EZA baited traps that attracted 94% of S. pictipes individuals, and 
since it is known that the presence of as little as 1% of the ZZA isomer as an impurity 
would significantly reduce the response of the moth to the lure (Tumlinson 1979), it can 
be concluded that the EZA lure used in this study was of high purity. The same scenario 
occurred with S. scitula: 80% of the individuals in the present study came to the EZA 
traps, while the other studies found the ZZA isomer to be the main lure. It would be in- 
teresting to find out if circumscribed and geographically isolated populations of the same 
species could be responsive to different pheromone isomers (there is some indication that 
attractancy to pheromones or mixtures may vary depending on what other species fly in 
the same area; Eichlin, pers. comm. ). 

In conclusion, with 24 species recorded here, southwestern West Virginia appears to 
have a rich sesiid fauna. Further collecting will doubtless add more species to the list 
(while this paper was being prepared, a male Synanthedan sigmoidea (Beutenmuller) was 
taken at Kanawha City in a 99:1 trap). The abundance data reported here should be inter- 
preted with caution: pheromone bait trap captures of males in specific locations may or 
may not reflect the overall abundance of particular species in the whole region. For in- 
stance, M. cucurbitae is certainly a common species in southwestern West Virginia where 
host plants are available, yet no individual was caught in any of the baited traps. It would 
be erroneous to conclude from this study that M. cucurbitae is a rare species. 


ACKNOWLEDGMENTS 
I thank Thomas Eichlin for identifying some of the more difficult species, and for offer- 
ing constructive criticism on the manuscript; Hermann Flaschka provided some of the 
more specific sex attractant samples; Claudia Thomas helped edit and prepare the paper. 
This paper was presented in part at the Annual Meeting of the West Virginia Entomologi- 
cal Society on 6 January 1995. 
LITERATURE CITED 


EICHLIN, T. D. & W. D. DuCKworTH. 1988. Sesioidea: Sesiidae. In Dominick, R. B. et 
al., The Moths of America North of Mexico. Fasc. 5.1. 


256 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ENGELHARDT, G. P. 1946. The North American clearwing moths of the family Aegeriidae. 
U.S. Natl. Mus. Bull. 190:146. 

NEAL, J. W., JR. & T. D. EICHLIN. 1983. Seasonal response of six male Sesiidae of woody 
ornamentals to clearwing borer (Lepidoptera: Sesiidae) lure. Environ. Entomol. 
12:206—209. 

NIELSEN, D. G., F. F. PURRINGTON & G. F. SHAMBAUGH. 1979. EAG and field responses 
of Sesiidae males to sex pheromones and related compounds. Proc. Symposium 
Pheromones of the Sesiidae, USDA, SEA, ARR-NE-6:11—26. 

SHARP, J. L. & T. D. EICHLIN. 1979. Distribution and seasonal occurence of Sesiidae (Lep- 
idoptera) attracted to E,Z and Z,Z acetate and alcohol. Proc. Symposium Pheromones 
of the Sesiidae, USDA, SEA, ARR-NE-6:35—46. 

SHARP, J. L., J. R. MCLAUGHLIN, J. JAMES, T. D. EICHLIN & J. H. TUMLINSON. 1978. Sea- 
sonal occurence of male Sesiidae in north central Florida determined with pheromone 
trapping methods. Florida Entomol. 61:245—250. 

SNOW, J. W., T. D. EICHLIN & J. H. TUMLINSON. 1985. Seasonal captures of clearwing 
moths (Sesiidae) in traps baited with various formulations of 3,13 octadecadieny] ac- 
etate and alchol. J. Agric. Entomol. 2:73—84. 

SOLOMON, J. D., F. L. OLIVErRA, J. H. TUMLINSON & R. E. DOOLITTLE. 1982. Occurence 
of clearwing borers (Sesiidae) in west central Mississippi. J. Georgia Entomol. 
WALID, 

TaFT, W. H. & J. W. SNow. 1991. A guide to the clearwing borers (Sesiidae) of the north 
central United States. North Central Regional Publication 394. 

TUMLINSON, J. H. 1979. The chemistry of Sesiidae pheromones. Proc. Symposium 
Pheromones of the Sesiidae, USDA, SEA, ARR-NE-6:1-10. 


VALERIU ALBU, 6 Kit Road, Charleston, West Virginia 25314, USA. 


Received for publication 5 October 1995; revised and accepted 18 April 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 256-263 


MIGRATORY ACTIVITY IN VANESSA CARDUI (NYMPHALIDAE) 
DURING 1992 IN WESTERN NORTH AMERICA, 
WITH SPECIAL REFERENCE TO EASTERN CALIFORNIA 


Additional key words: migration, population dynamics, weather, Owens Valley. 


Migrations of Vanessa cardui (L.) were unusually large in southwestern North America 
during 1991-1992, the largest since 1968 and 1973, providing a rare opportunity to study 
the complex mass behavior and population dynamics of this species (cf. Woodbury et al. 
1942, Abbott 1951). Here we summarize 1992 records for the region and present observa- 
tions made by one of us (DG) in Inyo County, California. The methods follow those out- 
lined in Giuliani and Shields (1995). Migration rates here (no./Smin/15m) are arbitrarily 
classified as light (1-29), small-scale (30—49), medium-scale (50—99) and large scale (>99). 
Vanessa cardui, like the monarch (Danaus plexippus L.), has a southward return migra- 
tion during the summer and fall (Emmel & Wobus 1966, Shapiro 1980, Myers 1985, Nel- 
son 1985, Giuliani & Shields 1995). 

Small numbers of migrating V. cardui were reported during February and March 1992 
in California, including: NW at Hemet (29 February, 12—22 March), NNW in San Diego 
County (9-10 March), NW-WNW near Bakersfield (11 March), NNE-WNW in Inyo 
County (15-17 March), and WSW in Ventura County (19 March) (pers. obs.; J. F. Emmel, 
in litt.; McKown 1993). A light migration was seen between Barstow and Yucca Valley on 
26 March (McKown 1993). Many northward migrators were seen near San Diego (27—30 
March) (R. Larson, pers. comm.), and there were several newspaper accounts of V. cardui 
plastering windshields during late March in the southern San Joaquin Valley. 


VOLUME 51, NUMBER 3 WOT 


One of us (DG) toured SE California and extreme W Arizona during 19—28 March to 
monitor V. cardui activities. Sightings increased southward from only a few non-migrators 
at NW San Bernardino County to a large-scale migration at Mecca, Riverside County; 
light to small-scale flights occurred in Imperial County and SW Arizona. Directions ob- 
served varied from NNE to W with most flights headed NNW-NW. 

On 28 March, R. E. Wells (in litt.) observed V. cardui in Baja California from Catavina 
to 5 km N Los Angeles Bay cutoff, where thousands of larvae were feeding on Malva L. 
(Malvaceae) with pupae utilizing the spines of young cardon cacti. Migration flights to the 
NNW were noted from Los Angeles Bay cutoff to 40 km SE Guerrero Negro. At San Lu- 
cas Cove during 28 March—6 April, adults flew NW—-NNW, including 11 km offshore. On 
9 April a large NNW-NW migration (3—30/km and over 300/min counted while driving) 
was seen in the vicinity of Guerrero Negro to ca. 65 km SE of there. Near Rosarito, up to 
93/km were counted and numbers increased near Punta Prieta on 9 April to an estimated 
620+ km, with larvae evident on the highway. 

Vanessa cardui significantly expanded its range northward during April. In Little Chino 
Valley (Yavapai County, Arizona), fresh adults migrated NW on 6-7 April in uniformly 
large numbers over at least a 16 km front; the main flight period lasted for 5 h each day 
(L. Muehlbach, in litt.). Migrations near Phoenix were completed by 10 April, with heavy 
NE movement in NE Arizona and NW New Mexico during 15—20 April even under cold, 
cloudy and snowy conditions; large sedentary populations were present during late April 
and early May in the North Rim, the Flagstaff area, and in the White Mountains (K. 
Roever, in litt.). A large flight passed through Denver, Colorado in an ENE-NE direction 
during 26 April—5 May (Scott 1992). Large numbers flew in the Los Angeles area in early 
April (M. & S. Foster, in litt.) and on 13 April in the western Mojave Desert (R. Larson, 
pers. comm.). There were nearly daily migrations in Inyo County throughout April, some 
large-scale in size, and migrations first appeared in Mariposa County after 5 April. Many 
migrating V. cardui were present by mid April in the Davis—Sacramento—Reno area, as 
well as in Salem, Oregon (R. Wescott, in litt.). On 27 April, large numbers flew SW in 
Owyhee County, Idaho (McKown 1992), and many faded migrators reached extreme SW 
British Columbia on 25—28 April (C. Guppy and R.P. Nelson, in litt.). From May to Au- 
gust, V. cardui became widespread in the United States and southern Canada (cf. Mc- 
Kown 1993, Swengel & Opler 1993). 

Inyo County observations. The first V. cardui appeared 3-17 March at Big Pine 
and displayed no apparent migratory behavior, as 3—6/day nectared on the blossoms of 
apricot trees. Light to large-scale migrations were seen 4—25 April at various Inyo County 
locations, with no migrations during 26—30 April. Beginning in May, flights became more 
complex as NW migrators intermingled with locally emerging adults that also flew north- 
ward. Occasional migrators appeared 1—3 May as local emergence began, soon producing 
densities comparable to those of April. Light to large-scale migrations were observed 4—17 
May, with a lull on the 13th. Few were seen during 18—21 May, but numbers increased in 
the following week (including two large-scale flights). Subsequently, few migrated on the 
Owens Valley floor, though light migrations continued at higher elevations in Inyo County 
until early October. 

Table 1 summarizes the numbers and directions taken by migration flights through 
Owens Valley during the spring of 1992. These flights reached maximum densities in mid- 
April and largely represent an influx from the SE, from the direction of the Sonoran 
Desert region of SE California, S Arizona and NW Mexico. Table 2 summarizes the large 
densities achieved by some of the migratory flights in the Inyo County region during April 
and May 1992. 

The final large-scale migration of V. cardui in Inyo County occurred on 25 May at 
1980—2315 m in the Inyo Mountains, with up to 152/5 min/15 m flying N—NE in the late 
morning. Migration rates were low below 2990 m 26—31 May, with almost no migrators at 
Big Pine after 1 June. Occasional migrators were observed at higher elevations through 
June, with small numbers also on flowers. 

Fig. 1 shows the shift in migratory flight directions during the summer. Southward mi- 
gration began in early July, overlapping with residual NW movement, and continued into 
October. These southward flights consisted of small numbers (<1/5 min/15 m) primarily at 


258 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Spring migrations of Vanessa cardui through Owens Valley, Inyo County, 
California in 1992. 


Date Maximum #/5 min/15 m Primary Geographic Directions 
March 17 1 NW-WNW 
April 1 2 NNW-—NW 
April 3 2 NW-WNW 
April 4 9 N-W 
April 5 59 variable (E—N—SW) 
April 7 30 NW-SW 
April 8 62 N-NW 
April 10 228 NW-—W 
April 14 ©) NNE-—NW 
April 15 81 NNW—-WNW 
April 16 265 WNW-WSW 
April 21 179 NW 
April 22 83 NW-WNW 
April 23 50 NNW-—W NW 


high elevations (less than 20% of the observations were below 3000 m). Only one migra- 
tion of 1992 showed a clearcut change of direction with time of day. On 5 April at Big 
Pine, 0800-1200 h, under clear skies and no wind, a medium-scale migration initially 
heading NW altered its direction counter-clockwise at a steady rate of 27°/h until heading 
SW as the sun moved clockwise across the sky. 

In Saline Valley at 480 m on 16 and 26 November, occasional individuals were seen in 
flight and on Baccharis flowers. No adults were seen in Saline Valley on 23-27 December 
under mostly sunny conditions with no wind (in some years a few have been seen there 
throughout the winter). Elsewhere, a few nectaring adults were observed 20—21 Septem- 
ber in the Providence and Dead Mountains, San Bernardino County, and several adults 
were seen on Lantana flowers at Hemet in November (J. F. Emmel, in litt.). On a trip to 
Baja California Norte in late November-early December, R. E. Wells (in litt.) observed 
only one adult at Miller’s Landing. 


TABLE 2. Inyo County, California large-scale migrations of Vanessa cardui, 1992. 

Date Locality Elev (m) Time (h) #/5 min/15 m Primary Directions 
April 10 —_ Lone Pine 1130 0830—0845 206-228 NW-—W 
Aprill6 8kmE Big Pine 1435 0920-0955 124-265 WNW-WSW 
April 16 Deep Springs Valley 1615 1300 128 SW-SE 
April17 Gilbert Summit 1890 1245 135 SE (some NW) 
April17 Gilbert Summit 1980 1400-1425 116-139 WNW 

(some SSE) 
April 20 N end Eureka Valley 1260 0645-0700 266—351 NW 
April 20 __ E of Big Pine 1830 1500 845 NW-—W NW 
April 21 White Mountains 2805 0815 104-105 NNE-NNW 
April 21 |= White Mountains 2650 0845 427 N—NW. 
April 21 Big Pine 1220 1145 179 NW 
May 6 Crater Mountain 1830 O0900—1100 >295 N 
May 10 Bridgeport, Mono Co 1970 0800 100 WNW-WSW 
May 12 White Mountains 2195 0745-0803 100—329 ENE-NNE 
May 17 N end Death Valley 1495 0830—0840 130-131 SE-ENE 
May 24 Inyo Mountains 2315 0945-1125 122-281 NNE-—NW 
May 25 Inyo Mountains AMIS) TS ey, NE-N 


VOLUME 31, NUMBER 3 259 


Ore Nise oe O) = Oo © 


© 10 20 0 40 60 6 70 8 © 100 110 120 130 140 150 160 170 180 180 200 210 220 200 240 250 260 270 280 200 300 310 320 330 040 350 060 


Fic. 1. Stacked bar chart showing change in flight direction from north to south of 
Vanessa cardui migrations above 2745 m in Inyo County, California. Black = May 12 to 
June 10, 1992; white = July 3-19, 1992. All directions are geographic: east = 0, north = 90, 
west = 180, south = 270. Vertical axis shows number of individuals observed within each 
10-degree interval. 


Mariposa County observations. V. cardui were first seen in Mariposa County mi- 
grating NNW-NW, on 6-§8 April near Mariposa, with a medium-scale migration flying 
NW on 9 April (1530-1645 h). On 19 April, NW migrating reached rates of 35—65/5 min/ 
15 m at 1035-1205 h, decreasing to occasional migrators on 20 April. A few dozen 
nectared at blossoms of Brodiaea Sm. (Amaryllidaceae) as well as yard Buddleja L. (Bud- 
dlejaceae) and apples. At Mariposa migration ceased whenever cloud cover appeared. 

At 1100 m at Jerseydale, V. cardui migrations did not appear until 21 April when a 
large-scale migration of wom individuals flew NNW at about 1400-1500 h. On 3-4 May 
occasional migrators occurred, along with resident V. cardui, that oviposited on lupines, 
Rumex L. (Polygonaceae), and other annuals. On 4 May at 1430 h, a small-scale migration 
of mostly fresh individuals flew NNE-—NNW,; flight was mainly to the NNE at 1530-1615 
h with numbers decreasing by 1645 h, and ceasing at 1743 h (occasional individuals flew 
in the opposite direction at 1726 and 1734 h). On 5 May at Jerseydale, migration began at 
0755 h in partly cloudy weather, with very few migrating at 0900-1000 h under overcast 
skies, with many nectaring on Chamaebatia foliolosa Benth. (Rosaceae). Only a few adults 
were seen after late May: on 9 June, 4 July, 25 July, and 11-13 October. On 11 June 
10-15% of the meadow thistles (Cirsium) had larval web-nests with dried frass of all sizes, 
but only four medium to large larvae were located (two alive, two dead). No larvae were 
found on lupines or other plants despite the earlier large migration numbers. 

At Jerseydale, six migrators were captured on 5 May and were placed at one location in 
the grass at the end of the day. These remained torpid overnight and began activity on 6 
May between 0640—0800 h by first rapidly vibrating their closed wings for about 2—5 min- 
utes; four of the six then flew in a complete circle before heading off erratically in a flight 
direction before usually landing again, while two others took off immediately without a cir- 
cle flight. The weather was overcast with dilute sunshine and calm conditions. Possibly the 
circling flight was performed for initial sun-orientation during overcast weather. In late 
April 1941 at 0650 h, six migrators that spent the night on a lawn flew up, circled around, 
and flew off to the NW (Abbott 1950:166). 

Migration times. No migrators were seen before sunrise. In Imperial County, as the 
sun was rising on 24 March at 0600 h, V. cardui adults were easily disturbed from the 
ground, with the first migrators observed at 0623 h; they appeared to first warm up on the 
ground at dawn, then went to flowers, then migrated. On 20 April in Inyo County, a single 
migrator was observed at 0555 h and another at 0600 h; at 0645 h a large-scale NW mi- 
gration abruptly appeared. Atop a 2315 m ridge in the Inyo Mountains on 24 May, counts 
were made of a large migration from 0500 to 1800 h. The first migrators were seen at 0545 
h and the last at 1735 h. The rate during successive 2-hour times from 0700 h was 14, 75, 
2:35, 52, 10; an estimated 517,000/km passed through the area on that day. 


260 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Cessation of daily migrations was determined in only a few instances. On 5 April a 
medium-scale migration at Big Pine continued until about 1705 h. On 9 April in the 
Owens Valley, occasional V. cardui were still migrating well after sunlight (until 1845 h) 
had left the tops of the surrounding mountains; four more were observed close together in 
flight against the moonlit clouds after 1900 h (D. Constans, pers. comm.). A medium-scale 
migration on 16 April ceased flying between 1725-1735 h. On 9 and 19 April at Mariposa, 
migration ceased at 1727-1730 h. On 9 May at Reno, Nevada, occasional migrators were 
last seen about 1800 h. 

Wind effects. On 22 April near Bishop a migration flew mainly NW during calm con- 
ditions, many 3—6 m high and a few up to 30 m high. Later, when strong gusty winds from 
the N were present, they all flew within 0.6 m of the ground, rising higher whenever the 
winds abated. Other reports have noted V. cardui migrations flying low to the ground 
against stiff or near gale-force winds (cf. Woodbury et al. 1942:172, Giuliani 1977). 

At 0900-1100 h on 6 May at an 1830 m summit south of Big Pine a large-scale migra- 
tion flew N off the top under a light S wind. Most were initially 1 m above the ground but 
continued flying horizontally or slightly upward as they poured over the summit, instead 
of descending down slope. Similar behavior in migrating V. cardui over hill and mountain 
summits has been reported elsewhere (cf. Wright 1906:37, Shields 1967: 112). On 23 May, 
atop a 2315 m ridge SE of Big Pine during a medium-scale migration with winds from the 
ESE-SSE, adults flew mainly NNE—NW up slope and continued upwards at the angle of 
the slope after passing the crest. 

When flying in a strong crosswind, individuals are often oriented to as much as 45° away 
from the observed direction of motion (toward wind direction). 

High fliers. Although most migrations stayed within 5 m of the ground, some high 
altitude flight was observed. On 23 March at Coachella, Riverside County, at 1200 h, a 
small-scale migration proceeding NW-—WNW flew 1.2 m to over 30 m high during warm, 
cloudy, breezy conditions. High-fliers proceeded in the same directions as those lower 
down. Flight numbers fluctuated dramatically, probably caused by the cooler cloud shad- 
ows encountered along their route. On 5 May near Big Pine, 0900-1100 h, under heavily 
clouded conditions, a light migration went N-NW; on four different occasions, individuals 
flying at under 2 m high climbed at a constant 20—40° angle until out of sight, while occa- 
sional migrators were seen fluttering or drifting at 15 m to over 30 m high. 

On 16-17 April at the SE base of a steep 700 m high ridge east of Big Pine, a large NW 
migration occurred under both heavy cloud cover and mostly blue sky conditions, extend- 
ing up to at least 300 m above the ground. Numbers observed by binocular reached as 
much as 600/min: many appeared to be drifting, hardly fluttering their wings, as though 
allowing themselves to be carried NW by the wind. Some of these drifters were spinning 
slowly (like drifting leaves) with no attempt to maintain a constant orientation. On the 
summit of this ridge, migrators came over the top no more than 15 m high. A similar case 
was noted on 24 May atop a 2315 m ridge SE of Big Pine when drifters were seen to over 
100 m high at 1715 h under a cloud shadow; they did not move their wings and often were 
rotated about so that they faced many different directions (including opposite to the di- 
rection of movement). High flying in V. cardui appears to differ from that observed in mi- 
grating D. plexippus (see Gibo 1986, Gibo & Pallett 1979) i.e., directed ascending flight 
vs. gliding in circles in rising air currents, although both exhibit some passive drift-gliding 
in tail winds aloft. 

Interactions. Occasional interactions by individual V. cardui on the ground toward 
migrators were observed. Seen most frequently in late afternoon or dusk and occasionally 
in the early morning, individuals would rise from the ground to follow or interact with a 
migrator and then return to the same spot. On 25 March in San Bernardino County, walk- 
ing at dusk caused adults to rise from the ground, including an occasional pets that flew up 
close together and then settled back down to the ground together. 

Pairs were often present in a migration, usually as one individual flying directly behind 
another. Seen from March through May, primarily during late afternoon or morming hours, 
they constituted as much as 10% of the total numbers in flight (over 50% on one occasion). 
The sex(es) of such pairs could not be determined, however. 

Courtship. Several courtships involving V. cardui were seen late in the day during 


VOLUME 51, NUMBER 3 261 


mid-April in Inyo County, though no actual matings were observed. On 11 April at 1615 h 
in strong winds, a pair descended to the ground and the male approached the female; the 
female avoided the male’s approaches, crawled into the base of a shrub, and the male 
perched on a twig 1.5 cm above her. On 17 April, after 1600 h, one pair landed in a shrub 
and each time the male approached, the female fluttered a few cm away. After the fourth 
try the male flew out and landed on the ground 1 m away. The female then immediately 
came out from inside the shrub and flew, sweeping low over the male, and the male rose 
and followed. Brown & Alcock (1990) reported four V. cardui pairs courting at 1605-1650 
h on a central Arizona hilltop; two mating pairs of V. cardui are recorded for 1800 h 
(Shields 1967, Brown & Alcock 1990); and two matings of the closely related V. kershawi 
(McCoy) occurred at 1730 h in Australia (Alcock & Gwynne 1988). 

Puddling. On 27 April, mostly very worn adults were fairly numerous on moist soil 
beside a creek. On 17 May in the Inyo Mountains during a large migration that progressed 
NNE, many were imbibing at muddy soils created by an irrigation ditch (D. Howell, pers. 
comm. ). Similarly, during a heavy migration in late May 1941 near Price, Utah, large num- 
bers congregated around roadside puddles (Know ton 1954). Puddling behavior is known 
to replenish sodium and water loss in male butterflies (Adler & Pearson 1982). 

Nectaring. V. cardui utilized many native desert flowers as nectar sources. In ap- 
proximate descending order of preference among the most readily used species were: 
Prunus andersonii Gray (Rosaceae), Dalea fremontii Torr. (Fabaceae), Salix L. (Sali- 
caceae), Amsinckia tessellata Gray (Boraginaceae), Encelia virginensis A. Nels. (Aster- 
aceae), Tetradymia DC. (Asteraceae), and Stanleya elata Jones (Brassicaceae). In the 
Owens Valley on 13 April an estimated 6000 adults were observed nectaring on a patch of 
P. andersonii 6 m in radius, and at Keeler on 10 April many thousands from a migration 
nectared on the numerous introduced Tamarix L. (Tamaricaceae) trees that were in full 
bloom. 

Oviposition, dwarf adults and hostplants. The large migrations of early April pro- 
duced many eggs throughout Inyo County, primarily on the abundant Amsinckia. During 
15-17 April, Amsinckia plants were found with up to 21 eggs per leaf along with small 
numbers of first instar larvae. By 19—21 April, many sites had more young larvae than ova. 
Small to dwarfed adult V. cardui were common in “May 1992, likely the result of their an- 
nual foodplants, such as Amsinckia and Cryptantha, drying out earlier than usual (i.e., 
contrast, migrating adults in Inyo County during April and after May were mostly re 4 
and large sized). At most sites in the Owens Valley, Amsinckia germinated during Decem- 
ber 1991 and had already advanced to the blooming stage by the time migrators arrived 
and oviposited in early April. By late April, the plants had mostly dried out and larvae were 
in instars 1—3 (few larvae or pupae could be found by 7 May). Millions of larvae likely per- 
ished there during late April due to the desiccation of the Amsinckia fields. 

On 18 April in the Chemehuevi Mountains, San Bernardino County, V. cardui larvae 
were abundant on Plantago insularis Eastw. (Plantaginaceae) with others on Amsinckia 
and Cryptantha, with fresh to worn adults common that were about 50% dwarfed (J. F. 
Emmel, in litt.). On 3-4 May, Emmel noted many nectaring adults near Amboy and in the 
Providence Mountains, San Bernardino County (also about 50% dwarfed). On 4 May at 
Jerseydale, Mariposa County, the junior author observed many small to dwarfed adults in 
a small-scale migration heading NNE in the late afternoon, most appearing fresh. Many 
adults of medium-small to dwarf size occurred in Inyo County from about 6—30 May. 
Dwarfs were common in Iron and Washington Counties, Utah, on 22—23 May, with a 
statewide dwarf population present in Colorado in late May (McKown 1993). 

V. cardui larvae were found in 1992 on the following hostplants that are additions to 
those reported for 1991 (Giuliani & Shields 1995): Boraginaceae: Cryptantha angustifolia 
(Torr.) Greene, C. circumscissa (H. & A.) Jtn., C. gracilis Osterh.; Fabaceae: Lupinus 
concinnus var. orcuttii (Wats) C.P. Sm., L. flavoculatus Heller, L. inyoensis Heller, L. mag- 
nificus Jones, L. pusillus var. intermontanus (Heller) C.P. Sm. A number of new hostplants 
were also recorded for the northern Central Valley of California in 1992 (Witham 1991). 

Crawling larvae. On 24 April at 670 m at the NW edge of Panamint Valley, V. car- 
dui larvae were abundant in an area where Amsinckia and Cryptantha plants were heavily 
eaten and desiccating. Many were crawling in straight lines on the ground in the morning, 


262 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


in roughly the same densities regardless of substrate and topography. Crawl speed aver- 
aged 1—2 m/min at densities up to 10/min over a | m line; two larvae crawled, respectively, 
11 and 18 m in 10 min and neither deviated more than 5° from a straight line. Though all 
possible crawl directions could be found, about 80% were going ENE—NNE and 20% 
WSW-SW with only a few in other directions. Larval measurements indicated that 
2nd—5th instars were involved in crawling (see Hammad & Raafat 1972) while many 4th 
and 5th instars remained on plants. While crawling, larvae often paused at plants they en- 
countered, investigated the leaves, and fed for varying lengths of time if it was a hostplant. 
If it was not a host, they continued crawling in their fixed directions. By 1000 h, with ris- 
ing ground temperatures, crawlers developed a strong tendency to leave their linear paths 
and head toward large objects (such as the observer), and most larvae soon ascended into 
Cryptantha and Amsinckia plants. However, many 3rd or 4th instars at this time were 
feeding on the leaves of Mentzelia, a plant they do not normally eat. 

Predators and parasites. In mid-April along the highway between Big Pine and 
Bishop, Inyo County, crows picked up dead and injured V. cardui adults that had been hit 
by cars (one had several in its beak). On 5 May at Big Pine, three english sparrows were 
pulling larvae off Amsinckia plants and eating them. On 24 April, black harvester ants 
were seen carrying off small, living larvae. On 5 May near Big Pine, over a dozen of a large 
unidentified wasp were searching Amsinckia plants and attacking any larvae they found. 
On 16 May at Owens Lake, a yellow-green crab spider, on a flower matching its color, 
had captured an adult (D. Constans, in litt.). On 2 July at 2745 m in the White Moun- 
tains, 11 small pupal cases of a wasp were found near where small larvae had been, in 
hidden and webbed locations between overlapping leaves of Cirsium drummondii T. & 
G. (Compositae). 


ACKNOWLEDGMENTS 


Richard A. Arnold, Lincoln P. Brower, David L. Gibo, Glenn A. Gorelick, James A. 
Scott, and Arthur M. Shapiro reviewed the manuscript al offered helpful esa eee 
Most of the plant identifications were made by Mary DeDecker. 


LITERATURE CITED 


ABBOTT, C. H. 1950. Twenty-five years of migration of the Painted Lady butterfly, Vanessa 
cardui, in southern California. Pan Pac. Entomol. 26:161—172. 

. 1951. A quantitative study of the migration of the Painted Lady butterfly, Vanessa 
cardui L. Ecology 32:155—-171. 

ADLER, P. H. & D. L. PEARSON. 1982. Why do male butterflies visit mud puddles? Canad. 
J. Zool. 60:322—325. 

ALCOCK, J. & D. GWYNNE. 1988. The mating system of Vanessa kershawi: males defend 
landmark territories as mate encounter sites. J. Res. Lepid. 26:116—124. 

Brown, W. D. & J. ALCOCK. 1990. Hilltopping by the Red Admiral butterfly: mate search- 
ing alongside congeners. J. Res. Lepid. 29:1—10. 

EMMEL, T. C. & R. A. Wosus. 1966. A southward migration of Vanessa cardui in late 
summer and fall, 1965. J. Lepid. Soc. 20:123—124. 

Gino, D. L. 1986. Flight strategies of migrating monarch butterflies (Danaus plexippus 
L.) in southern Ontario, pp. 172-184. In Danthanarayana, W. (ed.), Insect flight, dis- 
persal and migration. Springer-Verlag, Berlin. 

Gino, D. L. & M. F. PALLETT. 1979. Soaring flight of monarch butterflies, Danaus plexip- 
pus, during the late summer migration in southern Ontario. Can. J. Zool. 57:4393= 
1401. 

GIULIANI, D. 1977. Notes on the 1973 migration of Vanessa cardui. Pan Pac. Entomol. 
BOB. 

GIULIANI, D. & O. SHIELDS. 1995. Large-scale migrations of the Painted ae butterfly, 
Vanessa cardui, in Inyo County, California, during 1991. Bull. So. Calif. Acad. Sci. 
94(2):149-168. 

HAMMAD, S. M. & A. M. RaaFat. 1972. The biology of the Painted Lady Butterfly, Vanessa 
(Pyrameis) cardui L. Bull. Soc. Entomol. Egypte 56:15—20. 


VOLUME 51, NUMBER 3 263 


KNOWLTON, G. F. 1954. Migrations of Vanessa cardui, the Painted Lady butterfly, through 
Utah. Lepid. News 8:17—22. 

McKown, S. 1992. Painted Ladies migrate again. News Lepid. Soc. 4:71. 

. 1993. Season summary 1992. News Lepid. Soc. 2:25—49. 

Myers, M. T. 1985. A southward return migration of Painted Lady butterflies, Vanessa 
cardui, over southern Alberta in the fall of 1983, and biometeorological aspects of 
their outbreaks into North America and Europe. Canad. Field Nat. 99:147—155. 

NELSON, R. W. 1985. Southward migration of Painted Ladies in Alberta and British Co- 
lumbia. Blue Jay 43(1):7-15. 

ScoTT, J. A. 1992. Direction of spring migration of Vanessa cardui in Colorado. J. Res. 
Lepid. 31:16—23. 

SHAPIRO, A. M. 1980. Evidence for a return migration of Vanessa cardui in northern Cali- 
fornia. Pan Pac. Entomol. 56:319—322. 

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6:69—178. 

SWENGEL, A. B. & P. A. OPLER. 1993. Fourth of July butterfly counts, 1992 report. Xerces 
Society, Portland, Oregon. 75 pp. 

WiTHAM, C. W. 1991. The role of vernal pools in the 1992 mass dispersal of Vanessa car- 
dui with new larval hostplant records. J. Res. Lepid. 30:302—304. 

Wooppsuky, A. M., J. W. SUGDEN & C. GILLETTE. 1942. Notes on migrations of the 
Painted Lady butterfly in 1941. Pan Pac. Entomol. 18:165—176. 

WRIGHT, W. G. 1906. Butterflies of the west coast. Publ. by the author, San Bernardino. 
257 pp. 


DERHAM GIULIANI, P.O. Box 265, Big Pine, California 93513, USA, AND OAKLEY 
SHIELDS, 6506 Jerseydale Road, Mariposa, California 95338, USA. 


Received for publication 5 March 1995; revised and accepted 18 June 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 263-269 


DANCING WITH FIRE: ECOSYSTEM DYNAMICS, 
MANAGEMENT, AND THE KARNER BLUE 
(LYCAEIDES MELISSA SAMUELIS NABOKOV) (LYCAENIDAE) 


Additional key words: conservation, endangered species, metapopulation dynamics, 
sand and oak barrens, savanna, prescribed burning. 


The recent listing of the Karner Blue Butterfly (Lycaeides melissa samuelis Nabokov) 
as an endangered species (Clough 1992) has increased interest in managing and restoring 
populations of this charismatic invertebrate. The Karner Blue and other lepidopteran spe- 
cies are rapidly becoming symbols for restoring and conserving the barrens/savanna 
ecosystems that occur on well drained sand deposits in the Great Lakes Region and New 
England. The dynamic processes that produced unique botanical communities also pro- 
duced a highly specialized community of invertebrates adapted to this regime. Because of 
their general biological requirements, invertebrates are often closely linked to a few key 
ecological resources, such as specific soil types, edaphic conditions and/or individual host- 
plant species or genera (Panzer et al. 1995). 

The importance of oak barrens/savanna habitats to invertebrates is well illustrated by 
the Lepidoptera. In Ohio, the only midwestern state with a completed state-wide survey 
of all Lepidoptera species, the Oak Openings, Ohio’s only oak barrens/savanna community, 
supports the largest assemblage of imperiled butterflies and moths in the state. For exam- 
ple, five species of imperiled butterflies and 17 species of owlet moths (Noctuidae) occur 
in the Oak Openings, representing approximately 4% and 3% respectively, of the resident 
species in Ohio (Shuey et al 1987a, 1987b, Metzler & Lucas 1990, Iftner et al. 1992, Rings 
et al. 1992). The maintenance of this ecosystem is vital for the preservation of lepidopteran 


264 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Karner blue habitat: oak barrens in Newago County, Michigan. Note the sun- 
lit, grass-dominated opening surrounded by oak woodlands and numerous, fire-stunted 
oak and jackpine saplings within the clearing. 


biodiversity, as well as for other lesser known plants and animals in Ohio and the impor- 
tance of oak barrens/savanna communities to biodiversity maintenance in the other Great 
Lakes States is certainly similar to the situation in Ohio. For example, Panzer et al. (1995) 
list 17 species of butterflies that are primarily restricted to sand prairie, savanna and xeric 
prairie in the greater Chicago region. 

The decline of oak barrens/savanna lepidopteran communities can be attributed to sev- 
eral factors, but habitat loss, the disruption of ecosystem level processes and patch dynam- 
ics, and the collapse of metapopulation dynamics of many species are generally the pri- 
mary contributors. Here I discuss these intertangled processes, and the management 
implications and problems associated with each process as they relate to the Karner Blue 
(for ecological information regarding other imperiled midwestern lepidopteran species, 
see the species and habitat accounts in Rings et al. 1992 and Iftner et al. 1992). 

Habitat loss and fragmentation resulting from physical alteration. Habitat loss 
is often the most easily implicated factor contributing to the decline of most imperiled in- 
vertebrate species (Hafernik 1992) and the Karner Blue is no exception. To persist locally, 
Karner Blue populations require relatively large stands of the hostplant, blue lupine 
(Lupinus perennis L.) (Opler & Krizek 1984). Habitats supporting the butterfly are gener- 
ally open and sunny with scattered trees and shrubs (Fig. 1), and are dominated by grasses 
and other herbaceous species growing in well drained, sandy soils—in other words, 
healthy barrens/savanna communities (Zaremba & Pickering 1994). Oak barrens/savanna 
loss can be attributed to several factors, ranging from outright destruction to more subtle 
secondary impacts such as the encouragement of forest growth in areas of urban en- 
croachment. 

Oak barrens/savannas have been subject to the same trends that altered almost every 
ecosystem in eastern North America. The expansion of agriculture into new ecosystems 
was largely a process of trail and error: farming sand barrens was an error. In the trial pro- 
cess, many habitats were altered or destroyed and the local hydrology was often modified. 


VOLUME 51, NUMBER 3 265 


Fics. 2-3. Fragmentation of the dune and swale ecosystem (including dune-top oak 
barrens) of southern Lake Michigan. 2, the system in 1938. 3, the system as it appeared in 
1994. Note the fragmentation and almost complete olson of the remaining dune and 
swale fragments, surrounded by urban/industrial Gary and Hammond, Indiana. Scattered 
throughout this complex are habitats that support or have the potential to support the 
Karner blue. Fig. 2 courtesy of the Indiana Geological Survey. 


This trial process came to a halt during the prolonged drought of the 1930's, when it be- 
came apparent that the infertile soils of these communities could not support sustainable 
agricultural production. 

The unfortunate location of many regional barrens/savanna communities also con- 
tributed to their destruction, especially in New England. For example the Albany Pine 
Barrens sit adjacent to the city of Albany, New York, and the expansion of the city has, and 
still is contributing to the urbanization of this ecosystem (Dirig 1994). The Oak Openings 
ecosystem in Ohio is suffering the same fate as Toledo suburbs expand (Iftner et al. 1992, 
Grigore & Windis 1994); And the complex dune and swale communities which once lined 
southern Lake Michigan have been almost eliminated by industrialization and urbaniza- 
tion (Figs. 2 and 3). 

On a broader scale, the infertility of the sand soils themselves has led to the destruction 
of sand barrens communities. Many abandoned farms located in oak barrens/savanna 
ecosystems eventually reverted to federal and state ownership (via tax defaults), largely to 
become public forest land. Because the preservation of non-forest communities was not a 
high priority of national or state forests in the 1930's through the present, many oak bar- 
rens/savanna communities were converted into ‘productive’ use by conversion to pine 
plantations. These monocultures of stressed trees bear witness to the incomplete and 
short-sighted ecological planning of past eras. Degraded barrens communities continue to 
be primary targets for new developments such as industrial parks and residential commu- 
nities, possibly because the cost associated with acquiring barrens land is less than the cost 
for purchasing ‘productive’ agricultural lands. 

Habitat loss and fragmentation resulting from the disruption of ecosystem 
level processes and patch dynamics. Closely related to the impact of habitat loss is 
the elimination of ecosystem level processes. Oak barrens/savanna communities are 
among the most dynamic in the Midwest—the open habitats that support the Karner Blue 


266 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 
High 


Habitat 
Quality 


OAK Succession OAK Succession OAK 
i: Ae, ae >) VW >” 
aw BARRENS > D WOODLAND FF ,.¥ FOREST 


7 low / 
Tan 7 1 intensity 
endpoints 


intensity i 
lendeainis [ mienbence 
\ 


\ 
\ \ \ \ \ \ \ \ 

SS Sigs Ne ae ees Se as ie 
<——ee Disturbance(=Fire) <——& 


y, / / low ; 
high I intensity 
- : endpoints 
intensity l \ P { Disturbance 
endpoints Event 
\ \ 


Fic. 4. A simple model of the interaction between Karner blue habitat suitability, oak 
barrens succession, and fire disturbance. A: Optimal Karner blue habitat is early successional 
oak barrens; as succession proceeds, habitats become shaded and habitat quality decreases. 
B: Oak barrens, in the absence of disturbance, convert through succession to oak wood- 
land/oak forest communities. Note that while fire and other disturbances can re-set succes- 
sion to an earlier state, the exact outcome depends upon fire (disturbance) intensity and 
other mitigating factors. In the absence of disturbance, Karner blue habitat is eventually lost. 


were originally maintained by a steady procession of wildfires, which killed woody invasive 
plants while favoring fire-adapted dune and savanna communities. Without fire distur- 
bance, shade tolerant and fire sensitive species increase in density, and open barrens and 
savanna species decline. 

Functional oak barrens/savanna communities are in a constant but dynamic flux. Suc- 
cession pushes the community towards an association characterized by fire intolerant 
woody and shade tolerant herbaceous species, while fire disturbance realigns the commu- 
nity towards fire tolerant and shade intolerant species (Fig. 4). The original patch dynam- 
ics of these communities was in constant flux, and individual sites supported communities 
that reflected recent disturbance history. Although fire may have been a yearly occurrence 
within oak barrens/savanna ecosystems, the spatial distribution of the fire was less pre- 
dictable. For example, in the Albany Pine Barrens the point fire frequency may have 
ranged between 6 to 18 years, with a likely average frequency of once every 10 years 
(Givnish et al. 1988). Thus, these communities were composed of a constantly changing 
patchwork of habitats, reflecting the hit or miss nature of recent wildfires. Interdispersed 
through this patchwork were the recently disturbed sites supporting Karner Blue popula- 
tions. 

Unfortunately, our society tends to abhor wildfire because of the perceived destructive 
nature of fire. Thus, oak barrens/savanna ecosystems adjacent to urbanized areas are sub- 
ject to routine/reflexive fire suppression and state and national forests routinely suppress 
wildfires on their lands. With few positive attributes to associate with wildfire, active 
ecosystem management still remains controversial to the general public in many areas. 
Thus, society generally deprives these ecosystems of the very force that created them, a 
predictable and frequent fire disturbance regime. 

Urban and agricultural encroachment, in addition habitat elimination, fragment bar- 
rens/savanna communities by inserting non- or less-flammable land uses into a highly 


VOLUME 51, NUMBER 3 STE 


flammable ecosystem (Givnish et al. 1988). These barriers limit the occasional wildfire to 
small land tracts, reducing the potential for naturally spreading wildfire to maintain the 
ecosystem in an early successional state. In addition, urban encroachment increases the 
difficulty of using controlled burns to manage oak barrens/savanna communities because 
of the liability and perceived danger/nuisance to residents. 

Without the influence of a disturbance regime, oak barrens/savanna communities have 
succumbed to other community types. The impact of fire suppression on these communi- 
ties has been as great or greater than outright habitat destruction in most areas. For exam- 
ple, oak barrens are critically endangered and the Karner Blue is extirpated from Ohio’s 
Oak Openings, despite the “preservation” of over 9000 acres by state, local and private or- 
ganizations. Most of the habitats in the Oak Openings which once may have supported oak 
barrens/savanna have converted to young oak forest. Similarly, what remains of the Albany 
Pine Barrens in New York has converted into a largely overgrown ecosystem (Givnish et 
al. 1988). At its worst, land is dominated by black locust forests; at its best, dense scrub 
oak brushland is dominant. 

Disruption of metapopulation dynamics. The plants and animals that together 
form oak barrens/savanna communities are adapted to the ecosystem level processes 
which originally structured these communities. To persist regionally in this dynamic 
ecosystem type, invertebrates must cope with both the ecosystem patch dynamics as well 
as the forces driving patch dynamics. In simple terms, invertebrates populations must shift 
locations as quality habitats become available/unavailable and they must be able to survive 
wildfire, either directly or indirectly. While healthy metapopulations of the Karner Blue 
may seem to occupy entire barrens/savanna ecosystems, individual sub-populations are 
usually highly localized and isolated from neighboring populations by barriers of unsuit- 
able habitat. These isolated sub-populations are vulnerable to extinction from both com- 
munity succession and ecosystem disturbance regimes. 

Unfortunately, the Karner Blue is not well adapted to survive fire directly (e.g., Iftner 
et al. 1992, Swengel 1994). The very mechanism critical for creating and maintaining habi- 
tat for this species, fire, also kills all life stages of the butterfly (although there is emerging 
evidence that the Karner Blue may occasionally survive cool, low fuel-load fires, but re- 
quires better documentation). Recently burned habitats must be colonized or recolonized 
by individuals immigrating from nearby or adjacent habitats. Confounding this is the lim- 
ited dispersal abilities of the adults. Givnish et al. (1988) estimate that maximum dispersal 
distance for colonization of unoccupied habitats is approximately 0.5 miles. This agrees 
closely with values obtain in North Wales for the ecologically similar and related butterfly, 
Plebejus argus in North Wales (Thomas & Harrison 1992): i.e., metapopulation dynamics 
of P. argus over a seven year period indicated that the likelihood of colonizing suitable 
habitats decreased rapidly in habitats more than | km away from potential source popula- 
tions. These authors concluded that if the continuity of suitable habitat distribution was 
broken within an ecosystem, entire metapopulations of P. argus were likely to collapse. 

Because most oak barrens/savanna communities are suffering from the effects of fire 
suppression, optimal Karner Blue habitats are generally limited in size and widely dis- 
persed. This combination of reduced optimal habitat patch size combined with increased 
distance between optimal habitat patches has disrupted the metapopulation dynamics of 
the Karner Blue. For example, suitable but unoccupied habitats may not have a nearby 
Karner Blue source population from which colonization is possible. Likewise, occupied 
habitats may require recolonization following fires; recolonization has become less likely 
as the distance separating occupied habitats increases. In effect, the rate of localized pop- 
ulation extinction has been accelerated by declining habitat suitability and size, while the 
odds of new colonization events have declined as optimal habitats become increasingly 
fragmented due to succession and alteration. This disruption of metapopulation dynamics 
is currently causing the downward spiral of several metapopulations of the Karner Blue, 
even as regional attempts to restore these ecosystems proceed. 

The dance with fire. For the Kamer Blue, the interplay between habitat suitability, 
habitat distribution and patch dynamics, metapopulation dynamics and metapopulation 
persistence is complex. This is best illustrated by the historical distribution of the butterfly 
itself. The ecosystems known to support metapopulations of this butterfly are generally 


268 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


large, measured in tens of thousands of acres. Smaller sand barren/oak savanna complexes 
are less likely to have supported Karner Blues in historic times. This is probably a reflec- 
tion chance interplays between ecosystem processes and metapopulation dynamics: the 
larger the ecosystem, the better the odds that all the pieces fall together and populations 
persist. Smaller ecosystems may have provided fewer opportunities for population persis- 
tence, and Karner Blue populations did not persist to historic times. As Givnish et al. 
(1988) poetically state, persistent populations exist as a “flickering mosaic of Karner Blue 
populations, with some going extinct in a given area as others are being founded on sites 
recently burnt by colonists from sites burnt a somewhat longer time ago.” In smaller 
ecosystems, these populations may simply flicker out. 

However, given that almost every oak barrens/savanna community in the Midwest must 
now be actively managed to persist, much of the element of chance can be removed from 
Karner Blue management. With intensive management, which includes carefully planned 
burn units to create suitable habitats, Karner Blue populations should be manageable on 
preserves as small as 200 acres. At this scale, management would have to be almost me- 
chanical, with approximately 10-15% of the entire land-base burned annually, and the 
burn units configured to provide adequate dispersal opportunities for Karner Blues. 
Larger areas could be managed less mechanically, but would still require carefully planned 
management activities. Small management units could be used to establish core popula- 
tions within larger ecosystem management areas, from which dispersing butterflies could 
become more widely established. 

Finally, to protect against catastrophic disaster, several independent sets of Karner Blue 
populations should be maintained in each oak barrens/savanna ecosystem. Because of the 
flammable nature of the ecosystems, true wildfires that consume thousands of acres at one 
time are a reality. Because individual Karner Blue populations may succumb to such an 
event, independent core populations should be dispersed through the ecosystem to ensure 
that single catastrophic events cannot eliminate entire metapopulations. 

Preserve managers and stewards must struggle to re-establish the processes that cre- 
ated the barrens and savanna ecosystems they manage. If the Karner Blue is to survive, 
we must literally take it back to the big dance, where metapopulations swirled with patch 
dynamics to the music of fire. By managing remnant barrens and savanna communities in 
light of large-scale ecosystems processes, it should be possible to preserve not only the 
Karner Blue, but the untold other inconspicuous life-forms adapted to these ever chang- 
ing ecosystems. 


ACKNOWLEDGMENTS 


I thank the following groups and organizations for providing the access, opportunity 
and funding for my forays into Karner Blue habitats: The Nature Conservancy (Ohio 
Chapter), Indiana Department of Natural Resources (Division of Nature Preserves), Ohio 
Department of Natural Resources (Division of Natural Areas and Preserves and Division 
of Wildlife), Toledo Metro Park District, Michigan Department of Natural Resources, and 
most importantly, Save the Pine Bush, Incorporated (Albany, New York). Thomas Givnish, 
University of Wisconsin, provided valuable comments on an early draft of this manuscript 
and the thought processes of my fellow Karner Blue Recovery Team members have prob- 
ably been subliminally incorporated without attribution into portions of this manuscript. 


LITERATURE CITED 


CLOUGH, M. W. 1992. Endangered and threatened wildlife and plants: determination of 
endangered status for the Kammer Blue butterfly. Federal Register 57 (240):59236— 
59244. 

Diric, R. 1994. Historical notes on wild lupine and the Karner blue butterfly at the Al- 
bany Pine Bush, New York, pp 23-36. In D. A. Andow, R. J. Baker & C. P. Lane 
(eds.), Karner blue butterfly: symbol of a vanishing landscape. Misc. Publ. 84-1994, 
Minn. Agric. Exp. Stat., Univ. Minn., St. Paul. 

GIVNISH, T. J., E. S. MENGES & D. SCHWEITZER. 1988. Minimum area requirements for 
long-term conservation of the Albany Pine Bush and the Karner Blue butterfly: an as- 
sessment. Report submitted to the City of Albany, New York. 105 pp. 


VOLUME 51, NUMBER 3 269 


GRIGORE, M. T. & J. WINDIS. 1994. Decline of the Karner blue butterfly in the Oak Open- 
ings of northwest Ohio, pp 135-142. In D. A. Andow, R. J. Baker & C. P. Lane (eds. ), 
Karner blue butterfly: symbol of a vanishing landscape. Misc. Publ. 84-1994, Minn. 
Agric. Exp. Stat., Univ. Minn. St. Paul. 

HAFERNIK, J. E., JR. 1992. Threats to invertebrate biodiversity: implications for conserva- 
tion strategies, pp. 171-195. In P. L. Fiedler & S. K. Jain (eds.), Conservation biology: 
the theory and practice of nature conservation, preservation, and management. Chap- 
man and Hall, New York. 

IFTNER, D. C., J. A. SHUEY & J. V. CALHOUN. 1992. Butterflies and skippers of Ohio. Ohio 
Biol. Surv. Bull. New Series Vol. 9, No. 1. 212 pp. 

METZLER, E. H. & V. P. Lucas. 1990. An endangered moth in Ohio, with notes on other 
species of special concern (Lepidoptera: Saturniidae, Sphingiidae, Notodontidae, and 
Arctiidae). Ohio J. Sci. 90:33—40. 

OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins 
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PANZER, R, D. STILLWAUGH, R. GNAEDINGER & G. DERKOVITZ. 1995. Prevalence of rem- 
nant dependence among the prairie- and savanna-inhabiting insects of the Chicago 
Region. Natural Areas Journal 15:101—116. 

RINGS, R. W., E. H. METZLER, F. J. ARNOLD & D. H. Harris. 1992. The owlet moths of 
Ohio: Order Lepidoptera, Family Noctuiidae. Ohio Biol. Surv. Bull. New Series Vol. 
9, No. 2. 219 pp. 

SHUEY, J. A., J. V. CALHOUN & D. C. IFTNER. 1987. Butterflies that are endangered, 
threatened, and of special concern in Ohio. Ohio J. Sci. 87:98—106. 

SHUBY, J. A.. EH. MH. METZLER, D. C. IFTNER, J. V. CALHOUN, J. W. PEACOCK, R. A. 
WaArKINS, J. D. HOOPER & W. F. BABCOCK. 1987. Status and habitats of potentially en- 
dangered Lepidoptera in Ohio. J. Lepid. Soc. 41:1—12. 

SWENGEL, A. B. 1994. Observations on the effects of fire on Karner blue butterflies, pp. 
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THomas, C. D. & S. HARRISON. 1992. Spacial dynamics of a patchily distributed butterfly 
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ZAREMBA, R. E. & M. PICKERING. 1994. Lupine ecology and management in New York 
State, pp. 201-208. In D. A. Andow, R. J. Baker & C. P. Lane (eds.), Karner blue but- 
terfly: symbol of a vanishing landscape. Misc. Publ. 84-1994, Minn. Agric. Exp. Stat., 
Univ. Minn. St. Paul. 


JOHN A. SHUEY, The Nature Conservancy, Indiana Field Office, 1330 West 38th Street, 
Indianapolis, Indiana 46208, USA. 


Received for publication 22 August 1995; revised and accepted 13 July 1996. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 269-271 


LIFE HISTORY NOTES FOR THE PALLID EMPEROR MOTH, 
CIRINA FORDA (SATURNIIDAE) IN NIGERIA 


Additional key words: phenology, hostplants, Africa. 


Cirina forda Westwood has long been known as a serious pest of the sheanut tree, Vit- 
telaria paradoxa (Sapotaceae) in Nigeria (Golding 1929). Packard (1914) described the 
larva, and Boorman (1970) and Leleup and Beams (1969) provided brief accounts of the 
biology and phenology of this moth. Leleup and Deams (1969) reported Erythropheum 
africanum as a larval host in northern Zaire, but that the tree does not occur in Nigeria. 
The dried larvae of C. forda are referred to locally as “manimani,” and are of economic 


270 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


DEVELOPMENTAL STAGES 
Larva 


DJFMAMJJASOND J FMAMIJASONDJIEMAMNMIJI VV AS 


1988 1989 1990 (991 


Fic. 1. Phenology of Cirina forda in Nigeria, 1988-1991. 


importance as a food item among the Nupe tribe of Nigeria. For many decades, wild lar- 
vae have been collected, dried and sold to neighboring states within and outside the coun- 
try, and the insect is considered a delicacy (Ande 1991, Fasoranti & Ajiboye 1993). We re- 
port here on the life history and biology of C. forda in Nigeria, and its association with 
Vittelaria paradoxa. 

About 900 late instar C. forda were collected from V. paradoxa plants in August 1988, 
between km 107 and 108 on the Mokwa-Bida road in Niger State, Nigeria (9°05’N, 
5°59’E). Subsequently, rearings were conducted in the laboratory at the University of 
Ilorin between 1989 and 1991. Egg clusters laid in the laboratory were observed daily for 
hatching. Larvae from the same egg clutch were reared separately on cut and potted twigs 
of V. paradoxa and again on V. paradoxa plants outside in an ornamental garden. Twigs 
were replaced as necessary to maintain freshness, and larvae were transferred carefully by 
hand and placed on the mid rib or leaf margin of new foliage. A wooden cage (30 em x 30 
cm X 50 cm) with ten compartments was filled to a depth of 20 cm with soil. Ten larvae 
were then placed in each compartment. As soon as pupation began, the soil was in all com- 
partments examined for pre-pupae and pupae. Pre-pupa duration was defined as the pe- 


Fics. 2-6. 2, egg cluster. 3, pre-pupa. 4, fully formed pupa. 5, late instar. 6, adults (fe- 
male above, male below). 


VOLUME 51, NUMBER 3 Deal 


riod between soil penetration and actual pupation (in days). Subsequently, 30 pre-pupae 
were randomly selected and placed individually in soil 10 cm deep, each in cylindrical pa- 
per eclosion chambers (8 cm diam x 15 cm deep X 2 cm thick). The open end of the cylin- 
der was covered with a nylon mesh secured by a rubber band. Each compartment was ex- 
amined daily for emerging adults. Pupal duration was calculated as the mean number of 
days between pupal formation and date of adult emergence. Notes were also kept on adult 
longevity. 

Fig. 1 shows the phenology of C. forda for the period between 1988 and 1991. Adult 
moths lived for between 36 and 48 hours (mean = 39.7) and were found primarily in May, 
with peak oviposition at the end of the month. Figs. 2—6 show the immature stages of C. 
forda. The egg (Fig. 2) hatches after an incubation period of 30 to 34 days (mean = 31.8) 
into an active and voraciously feeding larva, and passes through 5 to 6 instars (Fig. 5) in 42 
to 50 days (mean = 47.5) between June and August. By the first few days in August, most 
of the larvae reach the pre-pupal stage (Fig. 3) and burrow into the soil. The pre-pupa de- 
velops into a pupa (Fig. 4) in 6—7 days and remains in diapause for 9 months (261 to 296 
days, mean = 267.5). Adult moths (Fig. 6) emerge in May of the following year. 

The life cycle of C. forda is tightly linked to the biology of its host, Vittelaria paradoxa. 
The only savannah species of the family Sapotaceae in Nigeria, V. paradoxa blossoms fully 
between May and August when mature fruits become available, and sheds leaves between 
November and February (Kaay et al. 1964). May and August is when the majority of C. 
forda larvae are developing in the field, and pupation takes place during the dry months of 
November and April. C. forda is univoltine in Nigeria and the phenology reported here 
agrees with those given by Golding (1929) and Boorman (1978). However, Leleup and 
Deams (1969) indicate that the active period for C. forda in Zaire is between June and 
September during dry months. In Nigeria, the active periods occur during the wet months 
of May and August. 


ACKNOWLEDGMENTS 


We acknowledge a postgraduate fellowship and Senate Research Grant awarded by the 
University of Ilorin, Horin, Nigeria. 


LITERATURE CITED 


ANDE, A. T. 1991. Some aspects of the biology of Cirina forda Westwood (Lepidoptera: 
Saturniidae) Unpubl. Ph. D. Thesis, Dept. Biol. Sciences, Univ. Ilorin, Nigeria. 327 


Senne J. P. T. 1970. The emperor moths (Saturniidae) of Nigeria. Nigerian Field 
35:99-122. 

. 1978. West African Butterflies and moths. Longman Group Ltd., London. 2nd 
Eid. 79 pp. 

FASORANTI, J. O. & D. O. AjiIBOYE. 1993. Some edible insects of Kwara State, Nigeria. 
Amer. Entomol. 93:113-—116. 

GOLDING, F. D. 1929. Preliminary notes on the pests of sheanut tree in Northern Nigeria. 
Bull. Dept. Agric. Nigeria 8:101—103. 

KEAY, R. W. J., ONOCHIE, C. F. A. & D. P. STANDFIELD. 1964. Nigerian trees. Dept. For- 
est Research. Ibadan, Nigeria. 495 pp. 

LELEvP, N & H. DaeEms. 1969. Les chenilles alimentaives du kwango canses de leur rar- 
efaction et mesures pre-conisees poury remedier. J. Agric. Trop. & Botany 
Appl. 16:1—21. 


A. TAIWO ANDE AND J. OLANIYAN FASORANTI Department of Biological Sciences, Uni- 
versity of Ilorin, P.M.B. 1515, Ilorin, Nigeria. 


Received for publication 6 December 1994; revised and accepted 26 January 1997. 


272 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Journal of the Lepidopterists’ Society 
51(3), 1997, 272-273 


SUPPLEMENT TO THE LIST OF THE BUTTERFLIES OF MICHIGAN 


Additional key words: Hesperiidae, Lycaenidae, Nymphalidae, faunal surveys, 
checklists. 


The following contribution represents an addition to the previously published faunal 
lists for Michigan Lepidoptera (Moore 1960, Perkins 1968, Nielsen 1970). Records range 
from 1941 to the present time, and represent collecting by myself. Two state zones (NLP 
= Northern Lower Peninsula; SLP = Southern Lower Peninsula) are recognized, and 
county names are given for individual records. A date with a trailing asterisk represents 
the earliest date captured whereas a leading asterisk represents the latest date of capture 
in a zone. Nomenclature follows Hodges (1983). 


Erynnis icelus (Scudder & Burgess) (Hesperiidae) SLP: Barry; *June 26 
Erynnis horatius (Scudder & Burgess) (Hesperiidae) SLP: Oakland 
Carterocephalus palaemon mandan (Edwards) (Hesperiidae) SLP Oakland; June 20* 
Thymelicus lineola (Ochsenheimer) (Hesperiidae) NLP: Newaygo. SLP: Huron 
Hesperia sassacus Harris (Hesperiidae) SLP: Barry 

Polites coras (Cramer) (Hesperiidae) SLP: Barry 

Wallengrenia egeremet (Scudder) (Hesperiidae) NLP: Newaygo 

Pompeius verna (Edwards) (Hesperiidae) SLP: *19 August 

Poanes hobomok (Harris) (Hesperiidae) SLP: Barry; *July 11 

Euphyes bimacula (Grote & Robinson) (Hesperiidae) NLP: Newaygo; *July 15 
Euphyes ruricola metacomet (Harris) (Hesperiidae) SLP: *August 19 

Papilio polyxenes asterius Stoll (Papilionidae) SLP: Huron 

Papilio troilus Linnaeus (Papilionidae) NLP: Newaygo 

Artogeia rapae (Linnaeus) (Pieridae) SLP: Lapeer, Sanilac, Tuscola ; 
Colias philodice Godart (Pieridae) NLP: Newaygo. SLP: Sanilac, Tuscola; May 4* 
Colias eurytheme Boisduval (Pieridae) SLP: Huron, Lapeer, Sanilac, Tuscola 
Lycaena phlaeas americana Harris (Lycaenidae) NLP: Newaygo. SLP: *October 1 
Hyllolycaena hyllus (Cramer) (Lycaenidae) SLP: Tuscola 

Epidemia dorcas (Kirby) (Lycaenidae) SLP: June 13, *August 19 

Epidemia helloides (Boisduval) (Lycaenidae) NLP: Newaygo 

Harkenclenus titus (Fabricius) (Lycaenidae) NLP: Newaygo 

Satyrium edwardsii ( Grote & Robinson) (Lycaenidae) SLP: *5 August 
Satyrium caryaevorum (McDunnough) (Lycaenidae) SLP: June 26* 

Satyrium liparops strigosum (Harris) (Lycaenidae) SLP: *August 19 

Everes comyntas (Godart) (Lycaenidae) NLP: Newaygo. SLP: Huron 
Celastrina ladon (Cramer) (Lycaenidae) NLP: Newaygo 

Aglais milberti (Godart) (Nymphalidae) SLP: Barry 

Vanessa virginiensis (Drury) (Nymphalidae) SLP: Barry 

Vanessa cardui (Linnaeus) (Nymphalidae) SLP: Huron 

Phyciodes tharos (Drury) (Nymphalidae) SLP: Lapeer; *Oct 1 

Euphydryas phaeton (Drury) (Nymphalidae) SLP: *August 7 

Basilarchia archippus (Cramer) (Nymphalidae) SLP: Tuscola 

Cercyonis pegala nephele (Kirby) (Satyridae) SLP: July 3* 


LITERATURE CITED 


HopcEs, R. W. (ed.). 1983. Check list of the Lepidoptera of America north of Mexico. E. 
W. Classey Ltd. and The Wedge Entomological Research Foundation. London. 284 


PP: 
Moore, S. 1960. A revised annotated list of the butterflies of Michigan. Occ. Papers Mus. 
Zool. Univ. Michigan 617:1—39. 


VOLUME 51, NUMBER 3 ATS} 


NIELSEN, M. C. 1970. New Michigan butterfly records. J. Lepid. Soc. 24:42—47. 
PERKINS, O. A. 1968. Addenda to the list of the butterflies of Michigan. J. Lepid. Soc. 
22:119—120. 


OWEN A. PERKINS, 2806 Linwood, Royal Oak, Michigan 48073-3023, USA. 


Received for publication 18 February 1997; revised and accepted 5 April 1997. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 273-275 


REDISCOVERY OF LETHE EUROPA TAMUNA WITH NOTES ON 
OTHER THREATENED BUTTERFLIES FROM THE ANDAMAN AND 
NICOBAR ISLANDS 


Additional key words: Legal protection, status reassessment. 


Drawing on data from the [UCN Conservation Monitoring Centre, the United Nations 
Environment Program (UNEP) (1987) listed six species of butterflies as threatened from 
the Indian coastal region. Four of these—one species and three subspecies—are endemic 
to the Andaman and Nicobar islands (Table 1). Three of these taxa have been termed 
“very rare’ while Graphium epaminondas Oberthur was termed “locally common” by both 
Evans (1932) and Ferrar (1948). Khatri (1996) recently reported that two of these taxa, 
Lethe europa tamuna de Niceville and Neptis sankara nar de Niceville, were extirpated 
on these islands. We present here new information on three of the four taxa rated “threat- 
ened” from the Andaman and Nicobar islands, including biological notes on the rediscov- 
ery of Lethe europa tamuna. 

Lethe europa tamuna de Niceville (Nymphalidae: Satyrinae). This is one of the rarest 
butterflies from the islands, being known previously from a single female specimen col- 
lected on Little Nicobar. Ferrar (1948) reported observing another female on Great Nico- 
bar some time before he left the islands in 1931. On a collecting trip to Great Nicobar Is- 
land in December 1996, the senior author observed four females, and found two eggs and 
two larvae of this butterfly at three localities in the Campbell Bay area of Great Nicobar. 
Both the adults and immatures were found along roadsides where the forests had been 
disturbed by human activity. One of the females was seen resting on moist sand on the 
banks of a stream. Another female was observed ovipositing on the upper surface of a leaf 
of the climbing bamboo, Dinochloa andamanica Kurz. Eggs were laid on leaves well 
within the clump, not on the fringes. The larvae (Fig. 1B) were sleeved and observed pe- 
riodically for about two weeks. They fed and passed through several instars, and confirmed 
D. andamanica as a host plant that supports development. We suspect this butterfly is not 
as rare as previously thought, but its status can be reliably assessed only after further stud- 
ies are conducted. 

Doleschallia bisaltide andamana Fruhstorfer (Nymphalidae: Nymphalinae). This 
butterfly has been considered rarer in the Nicobars (Car and Central Nicobar) than in the 
Andaman islands (Evans 1932, Ferrar 1948). Its cryptic habits have perhaps contributed 
to an underestimate of its abundance. We have observed eggs, larvae and adults of this 
butterfly at S. Andaman and at Great Nicobar. The larvae completed their life cycles on 
the plants on which they were found, when sleeved (see Table 1). We found from 15 to 58 
adults feeding on the small white blossoms of medium sized trees of Ligustrum glomera- 
tum Blume (Oleaceae) at Chidyatapu (Fig. 1D) in S. Andaman in 1994, 1995 and 1996. At 
Campbell Bay on Great Nicobar, 16 eggs were observed on P. album (Nees) Merr. (located 
in less than 30 min of search); 3 females were also observed in flight. 


274 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Threatened species of butterflies for the South Asian Seas region from the 
data base of the IUCN Conservation Monitoring Centre (UNEP 1987). Status represents 
ratings by Evans (1932) and Ferrar (1948); IUCN categories are as defined prior to Mace 
& Lande (1991) and Mace & Stuart (1994). All except Graphium epaminondas are listed 
as Schedule I species in the Indian Wildlife (Protection) Act, 1972 as revised to 1991. 


IUCN 
Taxon Region Distribution Status Category Larval Foodplant 
Lethe europa Dinochloa andamanica 
tamuna India SS. Nicobars very rare rare Kurz (Poaceae) 
Doleschallia S. Asia Andamans not rare rare Pseuderanthemum 
bisaltide album (Nees) 
andamana Merr. (Acanthaceae) 
Car Nicobars very rare Phaulopsis imbricata 
(Forst.) Sweet 
(Acanthaceae) 
Central Nicobars very rare 
Neptis sankara 
nar India N. Andaman very rare rare unknown 
Graphium 
epaminondas S. Asia Andamans not rare,  insuffi- Uvaria rufa Bl. 
locally ciently (Annonaceae) 


common known 


Graphium epaminondas Oberthur (Papilionidae: Papilioninae). Although this butter- 
fly was among the first collected from the islands (Hewitson 1874), nothing was known 
about its biology until its larval host plant and life cycle were worked out (Prashanth Mo- 
hanraj & Veenakumari 1994). It has a very short flight period with adults on the wing be- 
tween April and June (Prashanth Mohanraj & Veenakumari 1996). It is localized in its dis- 
tribution on S. Andaman. Mount Harriet National Park and some other localities on S. 
Andaman continue to support good populations of the butterfly, but we consider it has de- 
clined in abundance from the earlier reports by Evans (1932) and Ferrar (1948). The spe- 
cies is not currently threatened though some of its breeding localities have been destroyed 
(Prashanth Mohanraj & Veenakumari 1996). 

Neptis sankara nar de Niceville (Nymphalidae: Nymphalinae). About half a dozen 
historical specimens of this species have been collected from the Andaman islands (Ferrar 
1948). No specimens have been collected recently, and we have not encountered the spe- 
cies in our studies. If the butterfly is confined to N. Andaman (where Ferrar spotted his 
only specimen) then we may well have missed this species on our short, sporadic visits to 
the island. We doubt the species is extirpated, as N. Andaman is far less disturbed than 
S. Andaman. 

Over one fourth of the butterflies of the Andaman and Nicobar islands have been rated 
as “rare” or “very rare” by Ferrar (1948), the only person to have collected diligently for as 
long a period as eight years from these islands. Given the relative paucity of data, we feel 
that until more detailed field studies on Andaman and Nicobar butterflies are conducted, 
statements about the status of the islands’ species should be made with caution. 

We are grateful to A. K. Bandyopadhyay, Director, C.A.R.I., for supporting us in our 
studies on the natural history of the butterflies of these islands. We thank P. V. Sreekumar, 
Botanical Survey of India, Port Blair for the identification of the plant specimens, and 
Bikas Mondal, Tamil Das and Kinnu Ram for patient and cheerful assistance in the field. 
Permission to study insects from the National Parks and Sanctuaries of these islands was 
granted by the Chief Wildlife Warden, Andaman and Nicobar islands (Order No. 
CWLW/WL/47/1294). We also acknowledge financial assistance from the Ministry of En- 
vironment and Forest, Government of India, New Delhi (Order No. 14/44/91-MAB/RE). 


VOLUME 51, NUMBER 3 DATES 


Fic. 1. Lepidoptera from Andaman and Nicobar Islands: A, adult Lethe europa 
tamuna; B, larva of Lethe europa tamuna; C, the strikingly different larva of Lethe europa 
nudgara (ssp. tamuna is confined to the S. Nicobars while ssp. nudgara is restricted to the 
Andamans); D, Doleschallia bisaltide andamana feeding on a blossom of Ligustrum glom- 
eratum Blume. 


LITERATURE CITED 


EVANS, W. H. 1932. The identification of Indian butterflies (2nd revised Ed.). Bombay 
Natural History Society. Diocesan Press, Madras. pp. x + 454, xxii pls. 

FERRAR, M. L. 1948. The butterflies of the Andamans and Nicobars. J. Bombay nat. Hist. 
Soc. 47:470—491, 5 pls. 

HEwITsON, W. C. 1874. A list of butterflies, with descriptions of new species, from the An- 
daman islands. Ann. Mag. Nat. Hist. (4) xiv:356—358. 

KHATRI, T. C. 1996. Butterflies of the Andaman and Nicobar Islands: conservation con- 
cerns. J. Res. Lepid. 32:170—184. 

PRASHANTH MOHANRAJ & VEENAKUMARI, K. 1994. The larval food plant and life history 
of Graphium (Pathysa) epaminondas Oberthur—a papilionid endemic to the An- 
daman islands. Butterflies 7:24—34. 

. 1996. Host plants, phenologies and status of swallowtails (Papilionidae), Lepi- 
doptera, in the Andaman and Nicobar islands, Bay of Bengal, Indian Ocean. Biol. 
Conserv. 78:215—221. 

UNEP, 1987. Environmental problems of the South Asian Seas region: an overview. 
UNEP Regional Seas Reports and Studies, No. 82. United Nations Environment Pro- 
gramme. pp. li + 50. 


K. VEENAKUMARI AND PRASHANTH MOHANRAJ, Central Agricultural Research Institute, 
P. B. No. 181, Port Blair 744 101, Andamans, India 


Received for publication 16 April 1997; revised and accepted 21 July 1997. 


Journal of the Lepidopterists’ Society 
51(3), 1997, 276 


CORRECTION TO VOLUME 51 


In the article by Annette Aiello and Manuel A. Balcazar L., “The immature stages of 
Oxytenis modestia, with comments on the larvae of Asthenidia and Homoeopteryx (Sat- 


urniidae: Oxyteninae) which appeared in 51(2):105—118, the caption for Figure 1 contains 
an error. It should read Lot 85-126, not Lot 85-26. 


Date of Issue (Vol. 51, No. 3): 5 December 1997 


=: 


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CONTENTS 


MALE MATE-LOCATING BEHAVIOR AND YEARLY POPULATION CYCLES IN THE 
SNOUT BUTTERFLY, LiBYTHEANA BACHMANII (LIBYTHEIDAE) Ronald L. 
Rutowski, Barbara Terkanian, Ofer Eitan, and Andrea Knebel 


RELATEDNESS AND GREGARIOUSNESS IN THE ORANGE-STRIPED OAKWORM, 
ANISOTA SENATORIA (SATURNIIDAE) Adam H. Porter, Sean J. Cadaret, 
Scott A. Johnson, Hidetaka Mizohata, Andrea I. Benedetter, Cathleen 
L. Bester, Jennifer L. Borash, Scott D. Kelly, Gretchen S. Buehner, 
and Marilyn L. Sherman 


BIOLOGY OF THE BLACK-ANTENNA RACE OF PHYCIODES THAROS THAROS (NyYM- 
PHALIDAE) IN Ontario Paul M. Catling 


Two NEW SPECIES OF ASTERACEAE-FEEDING BuccuLaTRIX (BUCCULATRICIDAE) 
FROM CaLirorNiA Daniel Z. Rubinoff and Kendall H. Osborne 


A REVISION OF THE CERASTIS CORNUTA GROUP OF THE GENUS CERASTIS SUBGENUS 
Meratepsis (NoctuiwaE) Lars Crabo and J. Donald Lafontaine __. 


GENERAL NOTES 


Notes on the sesiid fauna of southwestern West Virginia Valeriu Albu 


Migratory activity in Vanessa cardui (Nymphalidae) during 1992 in western North 
America, with special reference to eastern California Derham Giuliani and 
Oakley Shields 


Dancing with fire: ecosystem dynamics, management, and the Karner Blue (Lycaeides 
melissa samuelis Nabokov) (Lycaenidae) John A. Shuey 


Life history notes for the Pallid Emperor moth, Cirina forda (Saturniidae) in Nigeria 
A. Taiwo ‘Ande and J. Olaniyan Fasoranti.2. 


Supplement to the list of the butterflies of Michigan Owen A. Perkins 


Rediscovery of Lethe europa tamuna with notes on other threatened butterflies 
from the Andaman and Nicobar islands K. Veenakumari and_ Prashanth 
Mohanray ics eT 


CORRECTION TO VOLUMEHD LT (2 So 


197% 


208 
218 
227 


237 


249 


256 
263 


269 
272, 


This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). 


1997 Number 4 


ISSN 0024-0966 


= ~—sdJourNAL 


9 ‘sn 3 of the 

_ Leprporterists’ Society 
} Be Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
4 uN Publié par LA SOCIETE DES LEPIDOPTERISTES 


- Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
a Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS 


EAA 


JAN 1 4 1996 


mt IO Dereniber 190% 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE CoUNCIL 


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Eric H. Merz.er, Immediate Past Mocens C. NIELSEN, 

President Vice President 
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Micuaet J. Smirn, Secretary 


Members at large: 


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Cover illustration: the Rosy Maple moth, Dryocampa rubicunda Fabr., a common saturniid found 
in eastern North America. Original pen and ink drawing by John Himmelman, 67 Schnoor Road, 
Killingworth, Connecticut, 06419, USA. 


TOUR NAL OF 
Toe LeEriporrTreERISTS’ SOCIETY 


Volume 51 1997 Number 4 


Journal of the Lepidopterists’ Society 
51(4), 1997, 277-287 


INTOXICATED LEPIDOPTERANS: HOW IS THEIR FITNESS 
AFFECTED, AND WHY DO THEY TIPPLE? 


WILLIAM E.. MILLER 


Entomology Department, University of Minnesota, 
Saint Paul, Minnesota 55108, USA 


ABSTRACT. Butterflies imbibing fluids at fallen, rotting fruits sometimes show signs 
of intoxication. Fallen fruits as well as woody plant sap-flows undergo natural fermenta- 
tion, which may result in frothy brews containing up to perhaps 3% ethanol. Many lepi- 
dopterans are attracted to volatile fermentation products, but studies of actual consump- 
tion are lacking. In laboratory choice tests, adults of Choristoneura fumiferana (Clemens) 
neither favored nor shunned 1% ethanol in plain or sweetened water for imbibing. Adults 
imbibing up to 1% ethanol were unimpaired in six of seven monitored fitness factors. One 
fitness factor, fertility, defined as the proportion of pairs reproducing, declined incremen- 
tally starting at concentrations of 0.5% ethanol. Two hypotheses are presented to explain 
lepidopteran intoxication in nature. 


Additional key words: Choristoneura fumiferana, Tortricidae, imbibing, fermenta- 
tion, ethanol. 


Butterflies may become sluggish and more easily captured while im- 
bibing fluids at fallen, rotting fruits, and collectors often use fermenting 
brews as baits (Norris 1936, Utrio & Eriksson 1977). Because the sweet 
fluids of fallen fruits and woody plant sap-flows may ferment (Janzen 
1977), it is assumed that lepidopterans imbibing them become intoxi- 
cated from fermentation products such as ethanol. Although the attrac- 
tancy of fermentation products to certain lepidopterans has been exper- 
imentally documented (Utrio & Eriksson 1977, Utrio 1983), studies of 
actual ferment consumption are lacking. Neither lepidopterans nor 
other insects are among the invertebrates featured in Winterstein’s 
(1919) classic treatise on narcosis. 

Unlike butterfly intoxication, moth intoxication does not seem to have 
been reported despite the fact that most of the experimental work on 
ferment attractancy utilized moths. Moths typically feed at dusk or after 
dark, times when they are difficult to observe in the wild. Also, in the 
era before sex lures, attractancy research was done to support prescrip- 
tions for trapping and monitoring lepidopteran pests in fruit orchards 


278 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(Dethier 1947, Green et al. 1960, Madsen & Morgan 1970), a context in 
which consumption and intoxication were irrelevant. Species in four 
moth families have nevertheless been recorded at natural sap flows, 
namely Noctuidae, Sphingidae, Geometridae, and Tortricidae (Norris 
1936, Foster & Tate 1966). 

Fermentation is the chemical alteration of carbohydrates by microor- 
ganisms. The microorganisms involved are usually yeast fungi and their 
close relatives, of which more than 500 species in 54 genera are recog- 
nized (Phaff et al. 1978). Different yeasts give rise to different fermen- 
tation products, and naturally produced ethanol is thought to be fairly 
common. It is not unusual for imbibed ferments to be described as 
frothy (Wilson 1926, Foster & Tate 1966, Simon & Enders 1978). Froth- 
iness results from the co-production of carbon dioxide with ethanol 
(Phaff et al. 1978). Yeasts are believed to be introduced by insects to 
fallen fruits and woody plant sap-flows (do Carmo-Sousa 1969, Phaff et 
al. 1978). Woody plant sap-flows result from wounding by a variety of bi- 
otic and physical agents, as well as from unknown causes (Wilson 1926, 
Ohman & Kessler 1964, Simon & Enders 1966, Radwan 1969). 

Here I examine whether or not individual C. fumiferana adults in the 
laboratory prefer diets for imbibing that contain 1% ethanol. I also com- 
pare the following seven fitness factors between groups of adults receiv- 
ing diets spiked with concentrations of 0.1—5% ethanol: fertility, life- 
span, preoviposition period, oviposition period, time to 80% oviposition, 
fecundity, and egg hatch. In designing and conducting this study, I drew 
heavily on previous personal experience with adult feeding in C. fu- 
miferana (Miller 1987, 1989). 


MATERIALS AND METHODS 


The adults used here were collected as pupae from balsam fir (Abies 
balsamea [L.] Mill.) and white spruce (Picea glauca |Moench] Voss) in 
two successive years at three sites within 6 km of Hovland, Cook Co., 
Minnesota. The pupae were sexed using the guide of Jennings and 
Houseweart (1978). Sexed pupae were placed one pair per container in 
1-pint (0.48 1) round cardboard ice cream cartons whose bottoms and 
tops were replaced with Petri dish bases and lids. Male and female pu- 
pae were matched developmentally so as to maximize eclosion syn- 
chrony. Pairs were assigned sequentially to different ethanol concentra- 
tion treatments within fitness experiments so that early and late eclosing 
pairs would be equally distributed throughout. A fresh sprig of balsam 
fir 5—8 cm long was placed in each pair container as an oviposition sub- 
strate. Containers were kept on a table in a laboratory maintained at 
25°C on a 121L.:12D fluorescent lighting schedule. Diets for imbibing 
were provided to moths by means of saturated 3-4 cc? synthetic 


VOLUME 51, NUMBER 4 279 


sponges. In the fitness experiments, there was one sponge per container, 
and the diet was renewed at intervals of 1—2 days. In the choice experi- 
ment, diets were provided only during tests. 

The choice experiment consisted of placing adults individually in a 
round I-gallon (3.8 1) 17.5 cm diameter cardboard ice cream carton with 
a glass cover, and observing each one for 20 min. Moth placement was 
at the center of the arena floor, 8 cm equidistant from two sponges on 
opposite sides of the floor, one soaked in a 1% solution of ethanol in ei- 
ther plain or sweetened water, the other soaked in the nonalcoholic 
equivalent. Sweetened water was 10% honey (v/v). If imbibing oc- 
curred, the time to its start was recorded. Each moth of each pair as- 
signed to the choice experiment was used once daily in a test near mid- 
day under regular laboratory lighting. 

In the fitness experiment with water-based diet, five concentrations 
of ethanol (0—5%) were provided, and in the experiment with 10% 
honey, three concentrations (0O—1%) (v/v) were provided. In both exper- 
iments, data were collected from pair containers once daily near mid- 
day. Records were made of female and male eclosion dates, number of 
eggs laid daily, and dates of male and female deaths. Foliage was in- 
spected for eggs with a 9 cm diameter reading glass. Eggs were removed 
from moth containers daily, counted under a stereomicroscope, and 
placed in labeled Petri dishes. Egg dishes were checked once daily to 
count the numbers of eggs hatching. Pairs were deemed fertile if the fe- 
male laid any viable eggs. At death, females were stored in a freezer un- 
til they could be dissected for counting unlaid mature eggs. Size and low 
stainability with methylene blue were the criteria by which unlaid eggs 
were deemed chorionated and thus mature or ripe (Miller 1987). 


RESULTS 


Diet-choice experiment. Thirteen female and 13 male adults 
were observed 92 times in the choice arena. Because choice results with 
plain and sweetened water diets were virtually identical, they were 
pooled. The pooled results show that no choice was made 43 out of 92 
times, an outcome consistent with previous findings that the moths do 
not always imbibe when given the opportunity (Miller 1989). Among the 
49 choices made, the chosen diet was as often nonalcoholic as alcoholic 
(Fig. 1). Adults choosing the nonalcoholic diet took 5.5 min (SD = 6.1 
min) on average to make a choice, while those choosing the alcoholic 
diet took 5.4 min (SD = 5.7 min). The moths that promptly made a 
choice walked directly to the sponge, often turning around once in place 
first. The preponderance of females over males in Fig. 1 is due to longer 
female lifespans. 

Fitness experiments. Moths receiving 5% ethanol became ex- 


280 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


No. 
choices 


50 
m Females Males 


40 


a5 Pos040 956 


30 27 


20 


10 


1% 0% 
ethanol alcohol Neither 


Fic. 1. Diet choices made within 20 min by individual Choristoneura fumiferana 
adults placed in the center of a 1-gallon (3.8 1) container with alcoholic and nonalcoholic 
diet sponges on opposite sides of the 17.5 cm diameter bottom. The alcoholic diet was 1% 
ethanol in either plain water or 10% honey water, and the nonalcoholic diet the same with- 
out ethanol. Results for the two experiments with different diet bases are pooled. The fre- 
quencies were tested for independence in a 2 x 2 contingency table using the G statistic. 


tremely intoxicated. They were lying on their backs within minutes after 
imbibing. Fourteen out of 16 did not reproduce, and although these re- 
mained alive for a few days, they always appeared comatose. At 1% 
ethanol, no signs of intoxication were evident. 

Fertility results from the two fitness experiments were pooled because 
of similar reproductive fractions in each experiment (Table 1). Fertility 
is broadly defined as the proportion of pairs reproducing (reproductive 
fraction). Pooled fertility dropped from 67% to 12% on diets of 0% to 
5% ethanol, respectively (Fig. 2). Fertility of 67% at 0% ethanol is typi- 
cal of normal laboratory fertility of C. fumiferana (Outram 1971, Miller 
1987, 1989). Distinct fertility reduction started at 0.5% ethanol. 

Results other than fertility from the two fitness experiments were not 
pooled because of the underlying differential effects of plain and sweet- 
ened water (Table 1; also Miller 1987, 1989). Within each experiment, 
preoviposition period, oviposition period, time to 80% oviposition, and 
lifespans of the sexes did not differ significantly among ethanol concen- 
trations 0O—-1% (Table 1). It must be emphasized that these results repre- 
sent only the fertile pairs at each ethanol concentration; results based on 


VOLUME 51, NUMBER 4 281 


TABLE 1. Performance of reproducing moth pairs receiving diets with different 
ethanol concentrations. Means are followed by SD’s in parentheses. Only reproductive 
fractions pooled from the two experiments differ significantly among ethanol concentra- 
tions (Fig. 4); differences in other factors among ethanol concentrations of 0-1% are not 
significant (P, > 0.20). Results for 5% ethanol were not included in the statistical analysis 
because of the small reproductive fraction. 


Ethanol concentration 


Factor 0% 0.1% 0.5% 1% 5% 
Water-based diet 
Reproductive fraction, pairs/pairs 12/16 WAG 10/16 10/17 2/16 
Mean preoviposition period (days) DES) (Os). AS (O70) 2G) BIL CLS) WOM) 
Mean oviposition period (days) VAL) DOC) SOCD) Siu) S55 (Os) 
Mean period female emergence to 
80% oviposition (days) CHUL) GO CLO) 63 CY) = OG (Sy Behe(OO) 
Mean lifespan (days) 
female 1O2AD) FRO TSC) — 1 (ZS), ESS (On) 
male Sod (DS) TOK) Gl Gis) Ge CUS) weal) 
10% honey-based diet 
Reproductive fraction, pairs/pairs 12/20 — 8/21 6/21 — 
Mean preoviposition period (days) 2.2 (0.6) = DOM (OS) Ds CLO) = 
Mean oviposition period (days) 13.6 (3.4) == WE (E59) IBS) _— 
Mean period female emergence to 
80% oviposition (days) 9.2 (1.9) = 8.8 (3.4) 9.8 (1.7) — 
Mean lifespan (days) 
female ismomecye>) == 16.4 (6.6) 19.8 (3.1) — 
male D4! (4) — 10.5 (4.9) 11.3 (6.2) a 


all pairs would differ moderately among ethanol concentrations. It is 
clear without statistical analysis that the two reproducing pairs receiving 
5% ethanol (Table 1) were severely impaired. These moths may have 
survived because of weak imbibing tendencies. 

Numbers of eggs produced, laid, and hatched showed no significant 
differences among concentrations of 0O-1% ethanol (Figs. 3, 4). Here 
again, the two pairs that reproduced on 5% ethanol clearly underper- 
formed (Fig. 3). The absence of increase in fecundity of females on al- 
coholic compared with the nonalcoholic diets suggests that alcohol was 
not metabolized for energy. 


DISCUSSION 


Technically, “intoxication” refers to behavior while “toxication” refers 
to deeper effects such as fitness. Throughout this paper I use the more 
familiar term to refer to both. Gomez's (1973) description of intoxication 
in a female Opsiphanes cassiae L. (Nymphalidae), quoted below, ap- 
pears to be the most detailed for a butterfly. The butterfly accidentally 
flew indoors and alighted near an uncorked bottle of wine containing 


282 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Percent pairs 
reproducing 
80 


P, < 0.001 
n= 144 
60 


40 


20 


0% 0.1% 0.5% 1% 5% 


Ethanol concentration in diet 


Fic. 2. Fertility or reproductive fraction of pairs of Choristoneura fumiferana adults 
receiving different ethanol concentrations for imbibing. Results for the two experiments 
with different diet bases are pooled. ; 


12% ethanol. Imbibing a drop of proffered wine, it began to act abnor- 
mally within 5 min. 

“First, some very slow up and down flapping of wings, followed by forewings being low- 
ered and directed forward with brisk movements several times, hindwings remaining up- 
right .. . [and moving] forward . . . until . . . propped far ahead of their normal resting po- 
sition. Antennae were lowered until they touched the table. . . . Movement of fore-, 
hindwings and antennae were repeated several times. . . . After a brief period of inactivity, 
a hopping spastic side-walking took place alternating with wing and antennae motions as 
well as a tremulous and agitated moving of the legs. More wine was offered to the insect 
which sipped it directly from my fingertip. . . . Another sequence of the behaviour de- 
scribed above was observed until all wings were placed flat on the table. . . . A few forward 
strokes of forewings followed by a very fast vibratory flapping preceded a period of inac- 
tion. A few minutes later the butterfly took flight in a close-spiralling pattern towards an 
incandescent light, hitting the hot bulb several times, alighting and again attempting flight 
to the light . . . close to which it finally perched. After a few hours it resumed normal be- 
haviour and flew away the next day.” : 


Where adults are short lived, as in Choristoneura fumiferana, intoxi- 
cation is more appropriately viewed in a fitness than behavioral context 
because reproduction is the predominant activity. The most ethanol- 
sensitive fitness factor proved to be fertility, broadly defined as the pro- 
portion of pairs reproducing (reproductive fraction). Fertility declined 
sharply from 49% at 0.5% ethanol to 12% at 5% ethanol (Fig. 2). It is 


VOLUME 51, NUMBER 4 283 


Mean no. eggs 
per female 


200 


Ripe £1 Laid MH Hatched 


2 
j 
Wy 
Y 
Y 
Y 
j 
Uj 


0% 0.1% 0.5% 1% 5% 
Ethanol concentration 


Fic. 3. Fecundity, oviposition, and egg hatch for Choristoneura fumiferana pairs re- 
ceiving different ethanol concentrations in plain water for imbibing. Results represent fer- 
tile pairs only. F-tests were done separately for each egg category. 


unclear at which interval fertilization was disrupted in the reproductive 
sequence of assembly, copulation, spermatophore transfer, sperm stor- 
age, and sperm use. However, the fact that females, and presumably 
males, are not known to imbibe until the third day of adulthood (Miller 
1989) suggests disruption following spermatophore transfer, an interval 
usually occurring on the first or second day of adulthood (Outram 1971). 
Nonfertile females were not dissected for spermatophores. 

No effects on C. fumiferana fitness were evident at 0.1% ethanol 
(Table 1, Figs. 2-4). However, it must be noted that actual quantities of 
ethanol ingested in this study are unknown. Previously, females were 
found to imbibe a mean of 4.5 mg of fluid per feeding (range of 
0.9—10.0 mg) (Miller 1989). 

One prominent instigator of sap-flows in forests inhabited by C. fu- 
miferana is the yellow-bellied sapsucker, Sphyrapicus v. varius (L.). 
This bird pecks squarish holes through the bark to the sap-conducting 
phloem in many species of woody dicotyledons and gymnosperms (Fos- 
ter & Tate 1966). Interiors of the holes are shaped so that sap collects in 
them. Sapsuckers and many other animals, including lepidopterans, 


284 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Mean no. eggs 
per female 


400 


Ripe Laid MH Hatched P->0.50 


300; 2 


200 


100 


0% 0.5% 1% 
Ethanol concentration 


Fic. 4. Fecundity, oviposition, and egg hatch for Choristoneura fumiferana pairs re- 
ceiving different ethanol concentrations in 10% honey for imbibing. Results represent fer- 
tile pairs only. F-tests were done separately for each egg category. 


feed at these reservoirs. Insects that visit them comprise much of the 
sapsucker diet (Foster & Tate 1966). Choristoneura fumiferana adults 
are also part of the sapsucker diet (Tate 1973), and presumably are 
among the tortricids at sapsucker feeding holes (Foster & Tate 1966). 
Intoxication signs sometimes noted in the birds themselves are attrib- 
uted to ingestion of fermentation products (Pearson 1936, Foster & Tate 
1966). Sugar concentration in the sap of feeding holes is typically 1-6%, 
sometimes higher (Tate 1973). Ethanol concentrations of ferments in 
nature apparently have not been measured, but typical sugar concentra- 
tions in sapsucker feeding holes would yield ethanol concentrations no 
higher than perhaps 3%, or one-half of the sugar concentration, assum- 
ing complete fermentation. However, fermentation may be self-limiting 
and thus incomplete (Jorgensen & Hansen 1948). Another source of fer- 
mentable sugars in forests is aphid honeydews, a food resource adult 
lepidopterans are also known to exploit (Pittioni 1923, Zoebelein 1956, 
Johnson & Stafford 1985). 

In certain Drosophila species, ethanol tolerance and metabolism are 
well developed and based on specific enzymes. Such species also exten- 


VOLUME 51, NUMBER 4 285 


sively exploit ethanol as a habitat cue (Chawla et al. 1981, Parsons 1981). 
By contrast, there is neither evidence for C. fumiferana adults metabo- 
lizing ethanol for energy, nor evidence for hormesis or small-dose en- 
hancement (Clarke 1990). Whether enzyme-based mechanisms of 
ethanol tolerance exist in any lepidopteran is unknown. 

As to why lepidopterans with imbibing capability tipple in the wild on 
ethanol or other fermentation products, I offer two hypotheses. 

First, lepidopterans that are neither attracted to nor repelled by fer- 
ments have life systems that lack the capability to use fermentation 
products as cues for finding adult feeding opportunities. Tiny amounts 
of fluids suffice for these small moths to imbibe, and their fitness is in- 
creased by water intake with or without dissolved sugars (Norris 1934, 
Kira et al. 1969, El-Sherif et al. 1979, Miller 1987, 1988). Lepidopterans 
in this group orient to the water at feeding sites, and intoxication is acci- 
dental, C. fumiferana being an example. 

Second, lepidopterans that are drawn to ferments have life systems 
with the capability to use fermentation products as cues for finding adult 
feeding opportunities. Members of this group are large bodied and re- 
quire ample nutrient fluids containing dissolved sugars and perhaps 
other ingredients (Portier & de Rorthays 1940, Lukefahr & Martin 
1964, Utrio 1983). If these lepidopterans become intoxicated from for- 
aging at ferments, the cost is tolerable given the food value of the unfer- 
mented fraction and possibly of the fermented fraction also. Opsiphanes 
cassiae, the intoxicated butterfly described earlier (Gomez 1977), is an 
example. Other examples include Nymphalidae in the fruit-and-sap- 
feeding subfamilies Brassolinae, Morphinae, Satyrinae, and Nymphali- 
nae (Young 1979), and many stout-bodied Noctuidae (Utrio 1983). 
Young (1979) has argued that eye spots on the wings of butterflies that 
forage at fallen fruits evolved as a defense to offset their greater vulner- 
ability to predation while on the ground, perhaps also while intoxicated. 

Future investigations toward understanding lepidopteran intoxication 
should include more species, more fermentation products, and the com- 
position of ferments at feeding sites. 


ACKNOWLEDGMENTS 


This work was supported by the Research Apprentice Program of the Cooperative State 
Research Service, U. S. Dept. of Agriculture, and by the Young Scholars Program of the 
College of Biological Sciences, University of Minnesota. I thank T. Do and D. Drekonja 
for laboratory assistance, and C. L. Boggs, J. E. Rawlins, A. M. Fallon, K. A. Mesce, and 
S. J. Weller for useful manuscript reviews. 


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Urrio, P. & K. ERIKSSON. 1977. Volatile fermentation products as attractants for 
Macrolepidoptera. Ann. Zool. Fenn. 14:98—104. 

WILSON, G. F. 1926. Insect visitors to sap-exudations of trees. Trans. Roy. Entomol. Soc. 
Lond. 74:243-—254. 

WINTERSTEIN, H. 1919. Die Narkose. Springer, Berlin. 319 pp. 

YOUNG, A. M. 1979. The evolution of eyespots in tropical butterflies in response to feed- 
ing on rotting fruit: an hypothesis. J. New York Entomol. Soc. 87:66—77. 

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Received for publication 13 February 1996; revised and accepted 9 December 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1997, 288-294 


HYDROXYDANAIDAL AND THE COURTSHIP OF 
HAPLOA (ARCTIIDAE) 


ROBERT B. DAVIDSON, CRYS BAKER, MELINDA MCELVEEN AND 
WILLIAM E.. CONNER 


Department of Biology, Wake Forest University, 
Winston-Salem, North Carolina 27109, USA 


ABSTRACT. Male Haploa clymene display large tubular coremata in the moments 
before mating. The coremata are deployed in the immediate vicinity of the female’s anten- 
nae and their eversion is a prerequisite for mating success. The major volatile component 
associated with the coremata of H. clymene and H. confusa is the dihydropyrrolizine hy- 
droxydanaidal. In H. clymene production of hydroxydanaidal is contingent on larval access 
to their natural hostplants containing pyrrolizidine alkaloid precursors. The widespread 
distribution of hydroxydanaidal in arctiid moths suggests a single early evolutionary origin 
for the ability to produce it. This origin appears to precede the divergence of three arctiid 
subfamilies: the Arctiinae, Pericopinae, and the Ctenuchinae. 


Additional key words: coremata, mating behavior, pyrrolizidine alkaloids. 


Male moths of the family Arctiidae possess some of the most morpho- 
logically elaborate scent-disseminating structures known in the Lepi- 
doptera (Birch et al. 1990). These structures are most often displayed in 
the moments before mating and are thought to play a role in sexual se- 
lection (Eisner & Meinwald 1995). The pheromonal signals they release 
have been characterized in several cases, and the compound hydroxy- 
danaidal (or the related compound danaidal) has been identified repeat- 
edly. Hydroxydanaidal is a volatile dihydropyrrolizine (Fig. 1) derived 
from defensive pyrrolizidine alkaloids (PAs) sequestered from the larval 
or adult food of each species (Conner et al. 1981, Boppré & Schneider 
1985, Krasnoff & Roelofs 1989). We here add Haploa clymene (Brown) 
and H. confusa (Lyman) to the growing list of hydroxydanaidal bearers, 
investigate the dependence of hydroxydanaidal production on larval diet 
in H. clymene, and describe the courtship of this species. We also sug- 
gest a single evolutionary origin for hydroxydanaidal production within 
the Arctiidae. 


MATERIALS AND METHODS 


Haploa clymene were collected as larvae on Eupatorium purpureum 
L. in Forsyth County, North Carolina. Adults were collected at black- 
lights and allowed to mate and lay eggs. Larvae were fed fresh leaves of 
E. purpureum, a member of a genus known to contain pyrrolizidine al- 
kaloids (Mattocks 1986), or Plantago rugelii Dcne., a plant known to be 
devoid of pyrrolizidine alkaloids (Cronquist 1981). Additional larvae 


VOLUME 51, NUMBER 4 289 


OH Ci® 


nf 


Fic. 1. R-(—)- hydroxydanaidal. 


were started on E. purpureum for their initial instar and then switched 
to an alkaloid-free Manduca artificial diet (Bell & Joachim 1976) until 
pupation. All larvae and adults were held at room temperature on a 
16L:8D photoperiod regime. Final-instar larvae of H. confusa were col- 
lected on Eupatorium maculatum L. and Lythrum salicaria L. by Scott 
Smedley in Ithaca, New York. These were fed E. purpureum until pu- 
pation. 

Coremata are tubular and inflatable scent-disseminating structures 
found in males of many species of arctiid moths. They were everted 
from between the 7th and 8th abdominal segments of 2—4 day old male 
Haploa by gently squeezing their abdomens. The coremata were excised 
with iridectomy scissors, and dropped into 100 ul of methylene chloride. 
Benzophenone was used as an internal standard. Quantitative chro- 
matography was carried out on a Hewlett Packard (HP) 55790A gas liq- 
uid chromatograph with an on-column injection port and flame ioniza- 
tion detector. Ultra I (methyl silicone) and Ultra II (phenyl methyl 
silicone) columns (Hewlett Packard; 25 m, 0.32 mm ID, 0.52 mm film 
thickness) were used. The carrier flow was 5.0m]/min with an initial 
temperature of 50°C held for 2 min; the temperature was then in- 
creased to 280°C at 5°C/min and held for 2 min. Authentic samples of 
R-(—)-hydroxydanaidal (99% pure) were provided by Jerrold Meinwald 
of the Department of Chemistry, Cornell University. 

Mass spectrometry and infrared spectroscopy were carried out on an 
HP 5890 GLC coupled with an HP 5965A infrared detector and an HP 
5970 mass selective detector. Compounds were separated on a 60 m, 
0.25 mm ID, 0.25 mm methy!] silicone column. The initial temperature 
was held at 100°C for 2 min and increased to 240°C at 2.5°C/min. 

Courtship behavior of H. clymene was observed and recorded in a lab- 
oratory wind tunnel (60 x 60 x 150 cm; windspeed = 25 cm/sec) under 
deep red illumination (<5 lux). Mating sequences were videotaped us- 
ing a BGC CCD-500E video camera and a JVC BR9000 video recorder. 


290 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Ultrasound was monitored using a QMC S200 bat detector and 
recorded on the audiotrack of the videotape. Some males were rendered 
incapable of everting their coremata by applying a cyanoacrylate ester- 
based glue to their partially everted coremata and allowing the coremata 
to retract. 


RESULTS 


Haploa clymene mate between 3 h and 7 h after the onset of sco- 
tophase. The courtship is initiated by the female through the release of 
a typical arctiid sex attractant blend. Female pheromone-releasing be- 
havior is readily apparent as the rhythmic exposure of the openings of 
tubular sex pheromone glands. The sex attractant, composed of Z,Z,Z- 
3,6,9-heneicosatriene and related compounds (Davidson 1995), stimu- 
lates males to fly upwind and seek females. When the male reaches the 
female he exposes a pair of inflatable cuticular tubes called coremata. 
They extend from their origin in the intersegmental membrane between 
the male’s seventh and eighth abdominal segments, often encircle the 
female’s abdomen, and curve together just above the head of the female, 
presumably stimulating her antennae with male courtship pheromone 
(Fig. 2). Genital contact is made and copulation ensues. Although Hap- 
loa have well-developed tymbal organs (Fullard & Fenton 1977, David- 
son 1995) like those that have been shown to be involved in the 
courtship of several arctiid species (Simmons & Conner 1996) their 
courtship is silent. 

The major volatile component associated with the coremata of both 
H. clymene (field collected as adults or fed E. purpureum through all 
their larval stages) and H. confusa (collected as final instar larvae on E. 
maculatum or L. salicaria and fed E. purpureum for the remainder of 
their larval life) matched an authentic sample of hydroxydanaidal in re- 
tention time on all three columns, IR spectrum, and mass spectrum (m/z 
(relative intensity): 151(3.5), 133(59), 104(100), 77(26), 51(35)). 

The coremata of male H. clymene raised through all larval instars on 
E. purpureum produced an average of 0.88 + 0.68 ug hydroxydanaidal/ 
individual (n = 13) with a range from 0.08 to 2.18 ug/individual. Males 
raised on Plantago rugelii (n = 1) or the alkaloid-free diet (n = 11) con- 
tained no detectable hydroxydanaidal in their coremata (detection limit 
0.01 g/individual). There were no apparent differences in the morphol- 
ogy of the coremata of H. clymene raised on the three diets. A compos- 
ite sample of the coremata of twenty H. confusa had a mean corematal 
titer roughly ten times lower (0.0685 wg/individual). 

The courtships of six unoperated male H. clymene (reared on E. pur- 
pureum), each paired with a female, were videotaped in the laboratory 
wind tunnel. In 5 of 6 cases the coremata of the males were clearly visi- 


VOLUME 51, NUMBER 4 291 


Fic. 2. Freeze-frame video sequence of the courtship of H. clymene. A, male approaches 
female (0.00 sec); B, male everts coremata (1.52 sec); C, coremata reach the antennae of 
the female (2.24 sec); D, pair in copulo (10.89 sec). Arrows mark everted coremata. 


ble and the courtship resulted in copulation. In the one case where the 
male did not evert his coremata, the courtship was unsuccessful because 
the female evaded the male by flying away. Three courtship sequences 
involving males rendered incapable of everting their coremata by gluing 
were all unsuccessful. In each case the female evaded the male and ef- 
fectively terminated the encounter. The difference between the success 
rates of unoperated males and the glued males was significant (Fisher 
exact probability, P = 0.047). Due to the small number of males avail- 
able sham-operated controls could not be performed for this experi- 
ment. However, previous studies indicate that the application of glue 
one segment forward on the abdominal venter had no effect on court- 
ship success in an arctiid (Conner 1987). 


DISCUSSION 


It is clear that Haploa clymene use hydroxydanaidal-laden coremata 
during courtship and are similar to Utetheisa ornatrix L. (Conner et al. 
1981), Pyrrarctia isabella (J. E. Smith), Phragmatobia fuliginosa (L.) 
(Krasnoff & Roelofs 1990), and Cisseps fulvicollis (Hubn.) (Meyer 1984) 


292 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


in exposing the coremata briefly just prior to contact between the male 
and female. Corematal exposure appears to be critical to courtship suc- 
cess; in its absence females evade males. The antennae of female H. cly- 
mene have been shown through electroantennogram bioassay to be sen- 
sitive to hydroxydanaidal (Davidson 1995), and thus it is likely that the 
courtship behavior of H. clymene is mediated by the hydroxydanaidal as- 
sociated with its coremata. The coremata of H. confusa also contain hy- 
droxydanaidal and, although courtship studies were not carried out on 
this species, they likely mate in a similar manner. The order of magni- 
tude difference in the hydroxydanaidal titer between the Haploa species 
is probably related to their dietary differences but further experiments 
will be necessary to verify this. 

Like Utetheisa (Conner et al. 1981), Creatonotus (Wunderer et al. 
1986), Estigmene, Pyrrharctia, and Phragmatobia (Krasnoff & Roelofs 
1989) production of hydroxydanaidal in Haploa is contingent upon dietary 
intake of PAs. Yet males reared on diets devoid of PAs have coremata that 
are visually indistiguishable from those of normal males, in striking con- 
trast to the morphogenetic effects of PAs on corematal development in 
Creatonotus (Boppré & Schneider 1985). The significance of the differ- 
ence in corematal development between these genera is not yet clear. 

Hydroxydanaidal usage is proving to be widespread within the Arcti- 
idae. Hydroxydanaidal (or the related dihydropyrrolizine danaidal) has 
been identified from the abdominal coremata of Estigmene acrea 
(Drury), Pyrrharctia isabella, and Phragmatobia fuliginosa (Krasnoff & 
Roelofs 1989); the genitalic coremata of Utetheisa ornatrix (Conner et 
al. 1981); the abdominal coremata of Creatonotus gangis (L.) and C. 
transiens (Wlk.) (Bell & Meinwald 1986); and the abdominal scent 
brushes of Paraeuchaetes pseudoinsulata Rego Barros (Schneider et al. 
1992) within the subfamily Arctiinae, and in the abdominal coremata of 
Cisseps fulvicollis (Krasnoff & Dussourd 1989) and the ventral valve of 
Cosmosoma myrodora Dyar (Ruth Boada, pers. comm.) within the sub- 
family Ctenuchinae. This broad phyletic distribution for hydroxy- 
danaidal usage in courtship suggests a single evolutionary origin within 
the Arctiidae. The common ancestor of the Arctiinae, the Pericopinae, 
and the Ctenuchinae appears to have been a PA-feeder (Jacobsen 1994; 
Susan Weller, pers. comm.) with, we propose, the ability to produce hy- 
droxydanaidal. We suggest that this single origin set the stage for the re- 
peated evolution of non-homologous scent-disseminating structures 
throughout the Arctiinae and the Ctenuchinae in a pattern consistent 
with sexual selection. Although the pericopine Gnophaela latipennis 
(Bdv.) has been shown to contain PAs (LEmpereur et al. 1989) no mem- 
bers of the subfamily Pericopinae have been studied with respect to hy- 
droxydanaidal usage in courtship. 


VOLUME 51, NUMBER 4 293 


ACKNOWLEDGMENTS 


We thank Scott Smedley for collecting H. confusa and Jerrold Meinwald for providing a 
sample of hydroxydanaidal. William Coleman and William Clapp provided GLC-IR-MS 
facilities. Becky Simmons and Ruth Boada assisted with the rearing of the insects and we 
thank Mindy Conner for editing the manuscript. This work was supported in part by a 
grant from Sigma Xi, The Scientific Research Society (R. B. Davidson), and the SURE 
program of the Department of Biology at Wake Forest University through NSF grant 
BIR-9321998 to Jerry Esch and Ron Dimock, Jr. 


LITERATURE CITED 


BELL, R. A. & F. G. JOACHIM. 1976. Techniques for rearing laboratory colonies of to- 
bacco hornworms and pink bollworms. Ann. Entomol. Soc. Amer. 69:365—373. 

BELL, T. W. & J. MEINWALD. 1986. Pheromones of two arctiid moths (Creatonotus tran- 
siens and C. gangis); chiral components from both sexes and achiral female compo- 
nents. J. Chem. Ecol. 12:385—409. 

BiRCH, M.C., G. M. Popry & BAKER, T. C. 1990. Scents and eversible scent structures of 
male moths. Ann. Rev. Entomol. 35:25—58. 

Boppre, M. & D. SCHNEIDER. 1985. Pyrrolizidine alkaloids quantitatively regulate both 
scent organ morphogenesis and pheromone biosynthesis in male Creatonotus moths 
(Lepidoptera: Arctiidae). J. Comp. Physiol. A 157:569—577. 

CONNER, W. E. 1987. Ultrasound: its role in the courtship of the arctiid moth Cycnia ten- 
era. Experientia 43:1029—1031. 

CONNER, W. E., T. EISNER, R. K. VANDER MEER, A. GUERRERO & MEINWALD, J. 1981. 
Precopulatory sexual interaction in an arctiid moth (Utetheisa ornatrix): role of 
pheromone derived from dietary alkaloids. Behav. Ecol. Sociobiol. 9:227—235. 

CRONQUIST, A. 1981. An integrated system of classification of flowering plants. Columbia 
Univ. Press, New York. 1262 pp. 

DavipsoNn, R. B. 1995. Courtship communication in Haploa clymene (Brown) and H. 
confusa (Lyman) (Lepidoptera: Arctiidae: Arctiinae): chemical characterization of 
male and female pheromones and description of the pheromone glands. Unpubl. M.S. 
thesis, Wake Forest University, Winston-Salem, North Carolina. 

DussourRD, D. E., C. A. HARVIS, J. MEINWALD & T. EISNER. 1991. Pheromonal adver- 
tisement of a nuptial gift by a male moth (Utetheisa ornatrix). Proc. Natl. Acad. Sci. 
(USA) 88:9224—9227. 

EISNER, T. & J. MEINWALD. 1995. The chemistry of sexual selection. Proc. Natl. Acad. 
Sci. (USA) 92:50—55. 

FULLARD, J. H. & M. B. FENTON. 1977. Acoustic and behavioural analyses of the sounds 
produced by some species of Nearctic Arctiidae (Lepidoptera). Can. J. Zool. 
Doo 224. 

JACOBSEN, N. L. 1994. Cladistic studies of the Arctiidae (Lepidoptera) and the genus 
Agylla (Arctiidae, Lithosiinae) using characters of the adult and larvae. Unpubl. Ph. 
D. dissertation, Cornell University, Ithaca, New York. 

KRASNOFF, S. B. & D. E. DussouRD. 1989. Dihydropyrrolizidine attractants for arctiid 
moths that visit plants containing pyrrolizidine alkaloids. J. Chem. Ecol. 15:47—60. 
KRASNOFF, S. B. & W. L. ROELOFS. 1989. Quantitative and qualitative effects of larval 
diet on male scent secretions of Estigmene acrea, Phragmatobia fuliginosa, and 

Pyrrharctia isabella (Lepidoptera: Arctiidae). J. Chem. Ecol. 15:1077—1093. 

. 1990. Evolutionary trends in the male pheromone systems of arctiid moths: evi- 
dence from studies of the courtship in Phragmatobia fuliginosa and Pyrrarctia isabella 
(Lepidoptera: Arctiidae). Zool. J. Linn. Soc. 99:319—338. 

LEMPEREUR, K. M., Y. L1& F. R. STERMITZ. 1989. Pyrrolizidine alkaloids from Hackelia 
californica and Gnophaela latipennis, an H. californica-hosted arctiid moth. J. Nat. 
Prod. 52:360—366. 

MATTOCKS, A. R. 1986. Chemistry and toxicology of pyrrolizidine alkaloids. Academic 
Press, London. 393 pp. 

MEYER, W. L. 1984. Sex pheromone chemistry and biology of some arctiid moths (Lepi- 


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doptera: Arctiidae): enantiomeric differences in pheromone perception. Unpubl. M.S. 
thesis, Cornell University, Ithaca, New York. 

SCHNEIDER, D., S. SCHULZ, R. KITTMANN & P. KANAGARATNAM. 1992. Pheromones and 
glandular structures of both sexes of the weed defoliator moth Paraeuchaetes 
pseudoinsulata Rego Barros (Lep., Arctiidae). J. Appl. Ent. 113:280—294. 

SIMMONS, R. B. & W. E. CONNER. 1997. Ultrasonic signals in the defense and courtship 
of Euchaetes egle Drury and E. bolteri Stretch (Lepidoptera: Arctiidae). J. Ins. Behav. 
9:909-919. 


Received for publication 9 July 1996; revised and accepted 30 October 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1997, 295-303 


WHY DO SOME MALE CALLOPHRYS XAMI (LYCAENIDAE) 
SHIFT THEIR TERRITORIES? 


CARLOS CORDERO 


Centro de Ecologia, Universidad Nacional Aut6noma de México, Apdo. Post. 70-275, 
Delegaci6n Coyoacan, D. F., México 


ABSTRACT. Ina Mexican population of the butterfly Callophrys xami at least 13% 
of the males defended two or more territories sequentially. There were two observed 
causes of territory shifts by males: aggressive displacement from their territories by other 
males (n = 2), and spontaneous shift to a different territory (n = 3); however, in 26 terri- 
tory shifts the causes were not determined. Evidence suggests that territories were in 
short supply during the study and, therefore, more territory shifts may have been the re- 
sult of aggressive displacement. The spontaneous shifts suggest that some males may move 
in search of a better territory after occupying one of low quality. 


Additional key words: behavioral variation, male competition, territoriality. 


In several butterfly species, males defend territories that are em- 
ployed exclusively for male display, mate location and courtship (Ru- 
towski 1991). Variation in territorial behavior in butterflies has been 
studied mainly in the context of alternative mate location strategies 
within a species (Davies 1978, Dennis 1982, Wickman 1985, 1988, Al- 
cock & O’Neill 1986, Dennis & Williams 1987, Alcock 1994), although 
some authors have also discussed the basis for differences between spe- 
cies in territorial vs. nonterritorial mating systems (Alcock 1985, Dennis 
& Shreeve 1988, Cordero & Soberé6n 1990, Wickman 1992). 

Although intraspecific variation in the number of territories sequen- 
tially defended by male butterflies has been documented (Alcock 1985, 
Knapton 1985, Alcock & O’Neill 1986), it has been specifically discussed 
in only one study (Robbins 1978). In some species, males spontaneously 
shift territory as a consequence of their normal migratory movements 
(Baker 1972). In non-migratory species there are at least two hypothe- 
ses to explain territory shifts; these hypotheses and some of their predic- 
tions are summarized in Table 1. 

In this paper, variation in the number of territories sequentially occu- 
pied by individual males of Callophrys xami Reakirt (Lycaenidae) is re- 
ported, and some of its possible causes and consequences are explored. 


MATERIALS AND METHODS 


The study was conducted in a 146.8 ha ecological preserve within the 
main campus of the Universidad Nacional Aut6noma de México, in 
Mexico City. This area is part of the Pedregal de San Angel, and is char- 
acterized by volcanic soil, rough topography, markedly seasonal rainfall, 
and xerophytic shrubby vegetation. 

Callophrys xami is a multivoltine butterfly that in the Pedregal de San 


296 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Two hypotheses to explain why males of non- migrant butterfly species might 
shift territories that they already occupy, and some predictions of these hypotheses. 


Hypothesis A: Males shift territories as a result of being aggressively displaced from 
their previous territories by intruder males. 


Prediction Al: Aggressive displacement of territorial males should be observable. 


Prediction A2: Successful territory holders (monoterritorial males) should be males with 
high resource holding power and, therefore, they should tend to be larger, more agile or 
more experienced than less successful territory holders (polyterritorial males). 


Prediction A3: Polyterritorial males, as a result of their displacement from high quality 
territories, should have a lower copulation success than monoterritorial males. 


Prediction A4: The incidence of territory shifts as a result of aggressive displacement 
should be higher when male density and, therefore, competition for territories is high. 


Hypothesis B: Males shift territories because they evaluate their current territories and 
voluntarily move in search of better ones. 


Prediction B1: Voluntary (spontaneous) territory shifts should be observable in territorial 
males. 


Prediction B2: Polyterritorial males should shift, on average, towards territories of higher 
quality (i.e., those with higher copulation rates). 


Prediction B3: Polyterritorial males, as a result of having spent some time in territories of 
poor quality, should have a lower copulation success than monoterritorial males. 


Prediction B4: Male density should be inversely correlated to the probability of finding an 
unoccupied territory of high quality, and therefore the cost of voluntary territory shift 
should be lower when density is low, and the probability of changing territory should be 
higher. 


Angel can be found at varying densities throughout the year (Sober6n et 
al. 1988). The population reaches peak density from October to January, 
although it is never abundant (Sober6n et al. 1988). The main larval 
food plant is the perennial Echeveria gibbiflora D. C. (Crassulaceae), 
which is abundant in the area (Soberon et al. 1988). Males are territorial 
and defend areas with well defined topographical limits, located beside 
or on natural or manmade trails; these areas lack concentrations of re- 
ceptive females and larval or adult food resources (Cordero & Soberén 
1990). Males actively defend their territories by means of different types 
of aggressive flights, for an average of five h per day (between 1000 and 
1500), and spend the rest of the time feeding and resting outside terri- 
tories (Cordero & Soberén 1990). Territories are occupied year after 
year and function as mate location and courtship stations (Cordero & 
Sober6n 1990, Cordero unpubl. data). Other details of courtshap behav- 
ior are given in Cordero (1993). 

A total of 159 territorial males was captured, individually marked on the 
wings with indelible felt-tip pens and their right forewing length measured 
with a caliper through the mesh of the net. Individuals were assigned to 
one of three wing-wear categories: 1 = similar to a recently emerged adult 


VOLUME 51, NUMBER 4 297 


(wings mostly green); 3 = very worn male (wings mostly brown with worn 
margins); and 2 = individuals intermediate between 1 and 3. 

Observations were made between 1 November and 20 December in 
1989, and between 10 November and 6 December in 1990. The num- 
ber of territories observed was 25 in 1989 and 19 in 1990; the number 
of days a territory was visited varied between 25 and 38 in 1989 and be- 
tween 14 and 24 in 1990. Observations were made in two ways: by walk- 
ing along transects joining groups of territories at least two times per 
day, on 31 days in 1989 and 11 in 1990, and observing each territory for 
a brief time; and by continuous observations through the daily territorial 
period in a group of occupied territories, during nine days in 1989 and 
13 days in 1990. 


RESULTS 


Most marked males were observed defending only one territory 
(86/99 males in 1989 and 52/60 in 1990; hereafter, monoterritorial males). 
Twenty-one males were observed sequentially occupying more than one 
territory (hereafter, polyterritorial males); these males represented 
13.2% of all marked males. Thirteen males occupied two territories, six 
males occupied three, and two males occupied four. Therefore, a total 
of 31 territory shifts was detected; however, the exact date of shifts was 
only determinable for 26 events. The median number of days polyterri- 
torial males occupied each territory was | (1.5 in fourth territory, n = 2); 
however, the range varied from <1 day to 14 days in their first territory 
(n = 20), to 1 to 2 days in their fourth territory (n = 2) (Table 2). Only 
one of the 55 marked males observed more than one day in 1983-1985 
occupied more than one territory, probably as a result of aggressive dis- 
placement (Cordero & Soberén 1990). Territories seem to be in short 
supply for the males of this butterfly, at least during peaks of male den- 
sity. In 14 of 17 cases, the site that a male had left was occupied by a dif- 
ferent male the same day or the day after. 

Direct support for Prediction Al (Table 1) was provided by two cases 
in 1989, in which the cause of territory shift clearly was aggressive dis- 
placement of the polyterritorial male by an intruder (for description of 
aggressive interactions see Cordero & Sober6én 1990). Two other cases 
in 1989 probably involved aggressive displacement and resulted in a 
territory shift. In the first case an aggressive interaction was observed af- 
ter which a male not previously in the territory began or continued de- 
fending it; less than an hour later, the male that had been defending this 
territory for the three previous days was observed defending a new ter- 
ritory. In the second case, a male was observed for over an hour defend- 
ing a territory, and then suddenly a different male was in residence; the 
first male was found defending a different territory 4.5 hours later. 


298 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


One way of testing Prediction A2 is by comparing the wing length (a 
measure of size and, possibly, resource holding power) and wing wear (a 
possible measure of age and experience) of males that are polyterritorial 
as a result of aggressive displacement with that of monoterritorial males; 
however, the small number of aggressive displacements observed in this 
study prevents statistical analysis. In one of three observations of aggres- 
sive displacement, the winning male was bigger (1.65 vs. 1.48 cm) and 
older (2 vs. 1), and in another it was smaller (1.55 vs. 1.62 cm) and 
younger (1 vs. 3) than the displaced male; data for the third case were 
not known. Of the two cases of probable aggressive displacement ob- 
served in 1989, the winning male was bigger in one (1.72 vs. 1.69 cm) 
and smaller (1.49 vs. 1.63 cm) in the other. These scant observations nei- 
ther support nor contradict Prediction A2. 

Since virtually all males observed during the course of this and previ- 
ous studies (since 1983) were territorial or were apparently trying to get 
a territory, the proportion of territories occupied in a given day was used 
as a measure of male density (Fig. 1). In 1989, the proportion of territo- 
ries occupied decreased through the study period (rg = —0.887, P < 
0.001, n = 35), but in 1990 no significant differences were observed in 
the proportion of territories occupied (rg = —0.305, P > 0.05, n = 17). 
Territory shifts were observed throughout the study periods in both 
years (Fig. 1). Contrary to Prediction A4, aggressive displacement was 
observed or suspected at both high and low densities in 1989. 

Regarding Hypothesis B (Table 1), we observed three cases of sponta- 
neous territory shifts (Prediction B1). In 1989, territorial male c moved 
from territory 3—4S to the contiguous territory 3—4N while inspecting a 
heterospecific butterfly, and perched in 3—4N without being detected 
by male b (who had been defending 3—4N since the previous day); after 
two minutes c aggressively displaced b and defended this “new” territory 
for the rest of that day as well as the next. No copulations were observed 
in territory 3—4S, in any of the eight days it was occupied by a male; four 
copulations were observed in territory 3—4N in the 23 days it was occu- 
pied by a male. Also in 1989, territorial male m moved spontaneously 
from territory V to territory IV (about 15 m away) aggressively displaced 
the previous resident and defended territory IV for one hour, returning 
afterwards to territory V. Male m occupied territory V four more days 
and later defended territory IV again on two days; this male was ob- 
served defending two other territories before defending territory V for 
the first time. One copulation was observed in the 31 days territory V 
was occupied; two copulations were observed in the 32 days territory IV 
was occupied. Finally, in 1990, territorial male 30a moved from territory 
E to territory F’ (which was unoccupied), about 25 meters away, and de- 
fended it for one day. This male was observed again defending territory 


VOLUME 51, NUMBER 4 299 


1989 


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Fic. 1. Proportion of territories occupied by males, territory shifts and matings ob- 
served during the study periods of 1989 and 1990. In 1989, only those days in which 17 or 
more territories were surveyed are included; in 1990, only those in which 13 or more ter- 
ritories were surveyed are included. Key: solid squares: observed aggressive displace- 
ments; empty squares: suspected aggressive displacements; diamonds: spontaneous terri- 
tory shifts; triangles: territory shifts due to unknown causes; solid circles: matings by 
monoterritorial males; sunbursts: matings by polyterritorial males; empty circles: propor- 
tion of territories occupied 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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VOLUME 51, NUMBER 4 301 


E on two days, four days after defending territory F’; afterwards he oc- 
cupied territory A for one day. No copulations were observed in any of 
the six and four days territories E and F’, respectively, were occupied. 
The fact that two spontaneous shifts were toward territories which ap- 
parently had higher copulation rates is in agreement with Prediction B2. 
The behavior of the last two males suggest sampling of territories, an 
idea implicit in Hypothesis B. 

In agreement with Prediction B4, the two spontaneous territory shifts 
witnessed in 1989 occurred when male density was low (Fig. 1). In both 
years, spontaneous shifts were observed in the second half of the study 
period and after most of the copulations were observed (Fig. 1), sug- 
gesting that a decreasing encounter rate with females may be used by 
males as a cue for voluntarily leaving the territory. 

Only two polyterritorial males were observed copulating, both in their 
second territory; these males were observed defending two territories 
and the causes of their territory shifts are unknown (one of these males 
was aggressively displaced from his second territory a few minutes after 
mating finished, and returned to his first territory). 


DISCUSSION 


In Callophrys xami some males shift territory because they are ag- 
gressively displaced from their territories by other males, or because 
they move spontaneously to a different territory. Given that the cause of 
84% of the territory shifts detected was unknown, the relative impor- 
tance of each of these causes cannot be determined. 

The direct observations of aggressive displacement indicate that com- 
petition for territories is an important cause of shifts between territories. 
Rapid re-occupation of abandoned territories also suggests intense com- 
petition for territories. Competition happens in spite of the availability 
of unoccupied territories (Fig. 1), suggesting that competition varies in 
space at a local scale, probably in response to limited male movement 
and differences in territory quality, and, temporarily, due to local 
changes in male density and territory quality. 

The existence of spontaneous territory shifts indicates that factors 
other than aggressiveness are responsible for some of the shifts. One 
possibility (Hypothesis B) is that males shift towards territories of higher 
quality (i.e., where mating rates are higher). We have insufficient data to 
test this possibility; however, the two observed copulations of polyterri- 
torial males occurred in their second territories. Furthermore, two 
spontaneous shifts were towards territories where copulation rates 
seemed to be higher. 

If the quality of prospective territories is difficult to determine for a 
male butterfly, males may simply tend to move to a different territory in 


302 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the hope of finding a better one. The time spent in a territory that is 
eventually abandoned may be necessary to determine its low quality or 
it may reflect a territory quality changing (decreasing) with time. Under 
these conditions we would expect to observe some cases of males shift- 
ing territory and returning to the previous one after some time, as was 
observed in two cases. Under this scenario, a smaller, and therefore 
more difficult to detect, difference between the average quality of pairs 
of territories sequentially occupied by males changing spontaneously 
might be expected. Intensive studies are needed to analyze the possible 
effects of territory characteristics on territory shifts. 


ACKNOWLEDGMENTS 


I thank C. Dominguez, W. G. Eberhard, C. Macias, J. Nufiez, J. Soberén, C. Wiklund and 
two anonymous referees for helpful comments. One anonymous reviewer and W. G. Eber- 
hard identified an error in a previous version of the manuscript. I particularly thank L. Gall 
for criticism and suggestions, as well as help improving the manuscript. Gabriela Jiménez 
and Rogelio Macias gave me valuable technical help in several phases of this research. I also 
thank my numerous field assistants who helped me observe butterflies. This research was 
supported by a Consejo Nacional de Ciencia y Tecnologia (México) scholarship. 


LITERATURE CITED 


ALCOCK, J. 1985. Hilltopping in the nymphalid butterfly Chlosyne californica (Lepi- 
doptera). Am. Midl. Nat. 113:69—75. 

. 1994. Alternative mate-locating tactics in Chlosyne californica (Lepidoptera, 
Nymphalidae). Ethology 97:103-118. 

ALCOCK, J. & K. O'NEILL. 1986. Density-dependent mating tactics in the gray hairstreak, 
Strymon melinus (Lepidoptera: Lycaenidae). J. Zool. (Lond.) 209:105—113. 

BAKER, R. R. 1972. Territorial behavior of the nymphalid butterflies, Aglais urticae (L.) 
and Inachis io (L.). J. Anim. Ecol. 41:453—469. 

CoRDERO, C. R. 1993. The courtship behavior of Callophrys xami (Lycaenidae). J. Res. 
Lepid. 32:99-106. 

CORDERO, C. R. & J. SOBERON. 1990. Non-resource based territoriality in males of the 
butterfly Xamia xami (Lepidoptera: Lycaenidae). J. Ins. Behav. 3:719—732. 

DaviEs, N. B. 1978. Territorial defense in the speckled wood butterfly (Pararge aegeria): 
the resident always wins. Anim. Behav. 26:138—147. 

DENNIS, R. L. H. 1982. Mate location strategies in the wall brown butterfly, Lasiommata 
megera (L.) (Lepidoptera: Satyridae): wait or seek? Entomol. Rec. J. Var. 94:209—214; 
95:7—10. 

DENNIS, R. L. H. & T. G. SHREEVE. 1988. Hostplant-habitat structure and the evolution 
of butterfly mate locating behaviour. Zool. J. Linn. Soc. 94:301—318. 

DENNIS, R. L. H. & W. R. WILLIAMS. 1987. Mate location behavior of the large skipper 
butterfly Ochlodes venata: flexible strategies and spatial components. J. Lepid. Soc. 
4]:45-—64. 

KNAPTON, R. W. 1985. Lek structure and territoriality in the chryxus arctic butterfly, 
Oeneis chryxus (Satyridae). Behav. Ecol. Sociobiol. 17:389—395. 

ROBBINS, R. K. 1978. Behavioral ecology and evolution of hairstreak butterflies (Lepi- 
doptera: Lycaenidae). Unpubl. Ph.D. dissertation, Tufts University, Medford, Massa- 
chusetts. 

RuTOWSKI, R. L. 1991. The evolution of male mate-locating behavior in butterflies. Am. 
Nat. 138:1121—1139. 

SOBERON, J., C. CORDERO, B. BENREY, P. PARLANGE, C. GARCIA-SAEZ & G. BERGES. 
1988. Patterns of oviposition by Sandia xami (Lepidoptera, Lycaenidae) in relation to 
food plant apparency. Ecol. Entomol. 13:71—79. 


VOLUME 51, NUMBER 4 303 


WICKMAN, P.-O. 1985. The influence of temperature on the territorial and mate locating 
behaviour of the small heath butterfly, Coenonympha pamphilus (L.) (Lepidoptera: 
Satyridae). Behav. Ecol. Sociobiol. 16:233—238. 

. 1988. Dynamics of mate searching behaviour in a hilltopping butterfly, Lasiom- 

mata megera (L.): the effects of weather and male density. Zool. J. Linn. Soc. 

Cesta 1 aOi Me 

. 1992. Mating systems of Coenonympha butterflies in relation to longevity. Evo- 

lution 44:141-148. 


Received for publication 17 January 1996; revised and accepted 27 August 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1997, 304-315 


EGG CANNIBALISM BY NEWLY HATCHED LARVAE OF THE 
SMALL WHITE BUTTERFLY, PIERIS RAPAE CRUCIVORA 
(PIERIDAE), ON AN ARTIFICIAL DIET 


MAMORU WATANABE AND TAKAKO OH’URA 
Department of Biology, Faculty of Education, Mie University, Tsu, Mie 514, Japan 


ABSTRACT. Newly hatched larvae of the small white butterfly, Pieris rapae cru- 
civora, wandered over an artificial diet without feeding for ca. 2 hrs after eating their own 
egg shells. When they encountered unhatched conspecific eggs, egg cannibalism occurred. 
Throughout the first instar, larvae fed on eggs and intermittently on the artificial diet. The 
duration of the first instar was significantly shorter for cannibals than for non-cannibals. As 
later first instars, the cannibals wandered randomly and only nibbled unhatched eggs. Egg 
cannibalism may help larvae exclude potential rivals from competing for nutrients when 
the host plant in the field is a limited resource. Because females lay eggs singly and sel- 
dom return to oviposit on the same host plant, siblicide in the field is presumably rare or 
absent. 


Additional key words: artificial diet, devouring, larval duration, nibbling, starved 
larvae. 


Females of the small white butterfly, Pieris rapae crucivora L., de- 
posit their eggs on the exposed leaves of cruciferous plants. Parasitic 
wasps, bugs, mites and ants have been recognized as major agents of egg 
mortality for pierid butterflies on cabbage (e.g., Harcourt 1966, Parker 
1970, Feltwell 1982) and on field cress, Rorippa indica Hiern (Yama- 
guchi & Watanabe 1993). Courtney (1986) pointed out that cannibalism 
is also a major cause of mortality in most Pierinae. 

In general, the newly hatched larvae of pierid butterflies eat their own 
egg shell before eating their host plants. However, the earliest hatched 
larvae often devour unhatched eggs on the same leaf (Rausher 1979, 
Watanabe & Yamaguchi 1993). Brower (1961) stated that eating egg 
shells may simply represent opportunistic egg cannibalism. Since egg 
cannibalism has been observed in high egg density, the behavior of lar- 
vae cannibalizing eggs has been regarded as abnormal (e.g., Feltwell 
1986, Warren 1992). However, cannibalism can strongly affect popula- 
tion density when resources are limited (e.g., Fox 1975), and show a 
density-dependent effect on population dynamics (Polis 1981, Elgar & 
Crespi 1992). The cannibalistic behavior of newly hatched larvae that 
occurs under crowded conditions in the absence of sufficient food has 
been reported in many species (e.g., Dempster 1983). When reared un- 
der crowded conditions, larvae of the orange tip, Anthocaris cardamines 
L., showed cannibalistic behavior (Feltwell, 1986). 

Egg cannibalism by larvae has been observed under conditions of nu- 
tritional deprivation in the laboratory (Hayes 1982). Stenseth (1985) 
concluded that cannibalism may evolve as the result of individual selec- 


VOLUME 51, NUMBER 4 305 


tion even in cases where food resources are not in extreme shortage. 
However, Shapiro (1981) found that egg cannibalism frequently oc- 
curred for P. protodice Boisd., in which a mechanism of avoiding ovipo- 
sition on the same leaf surface may have evolutionary implications. 
Watanabe & Yamaguchi (1993) found that intra- and inter-specific can- 
nibalism among pierid butterflies involved eggs and newly hatched lar- 
vae on the same leaf in the field. 

The present study was designed to provide insight into the mecha- 
nism of egg cannibalism by P. rapae under constant substrate conditions. 


MATERIALS AND METHODS 


Pieris rapae females were collected mainly in Nagano Prefecture, in 
the cool-temperate zone of Japan, during the summer of 1993. Mated 
females were obtained from the field and were allowed to deposit eggs 
on cabbage leaves. Eggs were laid during 2 h around noon on each sam- 
pling day. 

About 24 h after oviposition, each egg was placed on a medium con- 
taining artificial diet (Sato 1974) in a petri dish kept at room tempera- 
ture (ca. 25—-30°C). Wet filter paper approximately 8 cm in diameter was 
placed on the floor of each dish to reduce desiccation. Some eggs were 
placed on the wet filter paper for subsequent comparison of larval be- 
havior with that on the artificial diet. All of the eggs were in late devel- 
opmental stages, as identified by their egg color (yellowish orange). A 
detailed description of the developmental stages of eggs is given by 
Watanabe et al. (1993). 

Eggs offered to newly hatched larvae were derived from females 
placed on leaves on subsequent days. None of these eggs hatched ear- 
lier than the hatched larvae. Every egg was placed vertically on the food 
medium or filter paper like a naturally deposited egg. The arrangement 
of these eggs on the artificial diet in a petri dish is shown in Fig. 1. The 
number of eggs offered as food was 36 for each egg cannibalizing exper- 
iment. Twenty hatched larvae were tested. Since Watanabe & Yama- 
guchi (1993) found that the average distance between eggs deposited on 
a leaf of field cress, R. indica, was ca. 8 mm in the field, in this experi- 
ment all the eggs were placed 8 mm apart from each other. 

The position of each cannibalized egg was determined by counting 
the number of ‘steps’ from the previously cannibalized egg. A step rep- 
resents the space between eggs i.e., the distance measured in number 
of 8 mm units, because we did not know the actual distance of the route 
of the larva during 10 min observation intervals. For the first egg canni- 
balized, the location was the number of steps from the original point 
where the larva hatched. 

A stereoscopic microscope was used to observe each larva every 10 


306 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Petri dish (9cm) 


Fic. 1. Arrangement of eggs placed on artificial diet in 9 cm diam. petri dishes. Filled 
circle indicates an egg that will hatch first and become the cannibal. Squares, triangles and 
diamonds represent eggs 1, 2 and 3 steps away from the cannibal, respectively. 


min, from hatching to the first molting. Records were made of larvae 
feeding on their own egg shells, on the artificial diet, and on other eggs. 
Time spent moving between eggs and resting was also recorded. Larval 
position was recorded every 10 min, in relation to the position of the egg 
from which the larva hatched. 


RESULTS 


Immediately after hatching, all 20 larvae began to eat their own egg 
shells. This normally took about 90 min (Table 1). The larvae then 
moved out onto the artificial diet. 


VOLUME 51, NUMBER 4 307 


TABLE 1. Time elapsed for each feeding behavior from the time of hatching, for can- 
nibal and non-cannibal larvae of Pieris rapae. Values are minutes + SE, and are based on 
21 cannibal and 5 non-cannibal larvae. Asterisks denote significance at p < 0.05 for canni- 
bals vs non-cannibals. 


Feeding behavior Cannibals Non-cannibals 


end of feeding on own egg shell 97 + 11.1 88 + 18.5 
start of feeding on first egg 179 + 20.3 — 

start of feeding on artificial diet * 387 + 56.7 174 + 17.8 
duration of first instar * 3701 + 179.8 4863 + 133.8 


Larvae that were not presented with eggs (i.e., solitary larvae) began 
to feed on the artificial diet about three hours after hatching. The yel- 
low-brown mid-gut became greenish due to the color of the artificial 
diet. The larvae then wandered about on the diet, feeding intermit- 
tently, during the first instar stage. The duration of the first instar was 
about 82 h (=3.4 days). 

Newly hatched larvae that were presented with eggs began to eat the 
first egg three hours after hatching. This was not significantly different 
from the starting time of feeding on artificial diet by solitary larvae 
(=non-cannibals). The mid-gut remained yellowish brown for some time 
because they had not yet fed on the artificial diet. There was a delay be- 
fore the start of feeding on the artificial diet. The mid-gut of cannibal 
larvae did not become greenish until 6.5 h after hatching. This was sig- 
nificantly longer than the onset of greenish color in solitary larvae 
(P < 0.05 by F-test). Cannibal larvae wandered about with less feeding 
than solitary larvae. The duration of the first instar of the cannibals was 
significantly shorter (P < 0.05 by F-test). 

The newly hatched larvae fed on neighboring eggs (= 1 step) and then 
fed on eggs 2 steps away. As shown in Fig. 2, one of 20 larvae fed on 2 
eggs during the first instar stage, while another fed on 19 eggs (larvae 
fed on 8.3 + 4.7 eggs (SE) on average). Because the number of eggs of- 
fered as food was stable and they were evenly spaced, it can be seen 
from the data that, within the limitations of our method, cannibalism 
may not be a mortality factor in relation to density of unhatched eggs. 

Egg cannibalism first occurred 179 + 20 min after hatching (n = 14). 
All of the larvae ate eggs within one step. The second egg was eaten by 
the larvae 421 + 80 min after hatching (n = 15), and thereafter the third 
532 + 89 min (n = 13), the fourth 773 + 112 min (n = 13), and the fifth 
1108 + 204 min (n = 13). The time elapsed between each cannibalism 
event was thus 100—300 min. The tenth egg cannibalism was observed 
1618 + 95 min after hatching (n = 4). Larvae that were more than one 
day old displayed short intervals between cannibalism events. They 
moved out one step, and attacked adjacent eggs. The most active larva 


CTC 
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VOLUME 51, NUMBER 4 309 


WiWwy WY WiWvVW vo ov iy 


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30 


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Activity of newly hatched larva (mm/10min) 


0 10 20 30 40 50 60 70 80 90 


Time after hatching (hr) 


Fic. 3. Changes in movement by first instar larvae on artificial diet (+SD). Dots, open 
circles and small triangles represent cannibals, non-cannibals and starved larvae, respec- 
tively. Thick arrow shows average time required for larvae to eat their own egg shells. 
Each thin arrow indicates the average of each starting time for cannibals. Closed and open 
diamonds show the average starting times for feeding on artificial diet by cannibals and 
non-cannibals, respectively. Closed and open triangles indicate the average molting time 
to the second instar for cannibals and non-cannibals, respectively. Cross shows average 
time of death of starved larvae. 


taken for feeding on the diet and intermittent resting. Movement by lar- 
vae over 45 h old decreased to 5 mm/min. The older larvae often rested, 
and then molted into the second instar. The speed of movements of can- 
nibal larvae was generally similar to that of solitary larvae, but there 
were several high peaks of movement by cannibal larvae (Fig. 3). On av- 
erage, the cannibal larvae were more active than solitary larvae. 

Larvae hatched on the wet filter paper began to wander in the petri 
dish after eating their own egg shells. Their movement was the fastest. 
They were most active until 12 h after hatching. Their speed decreased 
gradually thereafter, and they became inactive after about 20 h. The av- 
erage longevity of the starved larvae was about 26 h (=1.1 days). 

The distance moved by larvae increased with time after hatching. As 
shown in Fig. 4, the change in the cumulative distance differed for can- 
nibal and solitary larvae (Kolomogorov-Smirnov test, 0.05 > P > 0.01). 
The cannibals moved further than the non-cannibals during the first 20 
h after hatching. However, both moved about 3000 mm during the first 
instar. A rapid increase in distance moved by the starved larvae was ob- 


310 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


5000 


1000 
500 


Cumulative distance moved (mm) 
oi 
[) 


0 10 20 30 40 50 60 70 80 90 


Time after hatching (hr) 


Fic. 4. Changes in the accumulated distance moved for first instar larvae on artificial 
diet. Dots, open circles and small triangles represent cannibals, non-cannibals and starved 
larvae, respectively. 


served on the wet filter paper. The cumulative distance moved differed 
significantly (Kolomogorov-Smirnov test) from the distances for canni- 
bal larvae (P < 0.01) and the larvae on artificial diet (P < 0.01). The 
starved larvae moved 1000 mm during their life span. 

The duration of feeding on a single egg also varied. It took a cannibal 
larva more than 20 min to consume an entire egg. Some larvae devoured 
the entire egg with intermittent resting, which increased the time re- 
quired for complete consumption. As shown in Fig. 5, there were a few 
larvae that finished eating the first egg within 20 min, but most spent 
much more than 20 min to consume the egg. Other larvae only nibbled 
parts of a victim egg shell before beginning to wander. Partial nibbling 
on an egg usually lasted less than 15 min. Fig. 5 also shows that most 
cannibal larvae did not spend much time to contact more than 10 eggs. 
There was a significant tendency for time spent cannibalizing to de- 
crease with the number of unhatched eggs encountered. Therefore, 
most cannibal larvae tended to devour entire eggs at the onset of feed- 
ing, and then nibble eggs later. All eggs that were nibbled did not de- 
velop further and did not produce larvae. 

Between the first to the fifth eggs cannibalized, larvae usually de- 
voured or nibbled an egg that was nearest (Fig. 6). The large average 
step number means that larvae moved long distances while feeding on 
eggs, and apparently sometimes ignored neighboring eggs while moving 
out randomly on the artificial diet. 


VOLUME 51, NUMBER 4 SL 


Number of larvae 


4G 500 
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Egg order cannibalized by a larva (E) 


Fic. 5. The relationship between the egg order cannibalized and the duration of can- 
nibalizing. 


DISCUSSION 


The present experiments show that larvae of P. rapae crucivora have a 
high propensity for egg cannibalism under laboratory conditions 
throughout the first instar. The newly hatched larvae wandered over the 
artificial diet without feeding, or on wet filter paper, for two hours after 
eating their own egg shells (newly hatched larvae also tend to wander 
actively on host plant leaves in the field i.e., they seem to search for eggs 
on the leaves similar to their behavior on the artificial medium in this ex- 
periment; Watanabe, unpubl. data). While wandering on artificial 
medium, larvae attack eggs that are encountered. If the larvae cannot 
find eggs, they begin to feed on the artificial diet. Watanabe and Yama- 
guchi (1993) found that larvae on leaves with conspecific eggs behaved 
as cannibals before starting to eat leaves in the field. 

Larvae did not feed on the artificial food medium for two hours after 
hatching, but wandered. They may waste energy during this period. 
However, starved larvae were able to wander actively on wet filter paper 
for 12 h. Therefore, newly hatched larvae may not be adversely affected 
by failing to feed during their first two hours, as this time may be for lo- 
cating eggs in the field. 

Two kinds of egg cannibalism were seen: consumption of the entire 
egg, and nibbling of part of the egg shell. Both kinds of cannibalism 
were fatal to unhatched larvae. The former presumably provides the 
cannibals with nutrients, whereas nibbling may exclude unhatched con- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


312 


(=) 
(90) 


>11 eggs 


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Steps required to reach the cannibalized egg 


FIG. 6. 


distribution. See 


-number of eggs cannibalized. 1-5, 6-10, 


Frequency distributions of the step 
>11 eggs mean the order of eggs cannibalized. Dots show the Poisson 


text for further elaboration. 


VOLUME 51, NUMBER 4 313 


specifics that would otherwise be competing for food. Therefore, possi- 
ble advantages for the cannibal include both nutrient gain and the elim- 
ination of competitors (Baur & Baur 1986). 

Eggs of -P rapae crucivora contain amino acids and organic com- 
pounds that are not directly derived from leaves (Porter 1992). Such 
materials might facilitate larval development during the first instar. Can- 
nibalism increases the growth rate of larvae that eat eggs (e.g., Dickin- 
son 1992, Agarwala & Dixson 1992). Osawa (1992) stated that first instar 
larvae of the lady beetle, Harmonia axyridis Pallas, developed faster 
after eating conspecific eggs. In P. rapae crucivora, however, there were 
no significant differences between cannibalizing and non-cannibalizing 
larvae over the whole larval period with regard to adult size and weight, 
or female fecundity (Watanabe, unpubl. data). We observed no egg can- 
nibalism by second, third, fourth and fifth instar larvae, despite Ya- 
mamoto’s (1981) finding that the eggs of P. rapae crucivora and P. napi 
L. were eaten by older larvae. 

Courtney & Courtney (1982) stated that cannibalism is concentrated 
upon particular host individuals, because of contagious egg distribu- 
tions. Brower (1961) stated that egg cannibalism is density-dependent 
in the case of the Queen butterfly, Danaus gilippus. Polis (1981) noted 
that cannibalism can also be a tactic to gain exclusive use of resources 
that serve as both food and habitat. For larvae, there are advantages in 
being single: moye food, and less chance that other members of the 
same family group will become parasitised or eaten at the same time 
(Feltwell 1986). The proportion of eggs surviving was a function of lar- 
val density in an Australian population of P. rapae (Jones & Ives 1979). 
Watanabe & Yamaguchi (1993) observed that, as a rule, a single larva 
settled on a single leaf of the field cress, R. indica, suggesting that newly 
hatched larvae may have consumed unhatched eggs on the same leaf. 

Egg cannibalism in P. rapae crucivora may be advantageous for larval 
survival on limited resources as well as for the intake of nutrients. While 
devoured eggs were usually those nearest to the larva, nibbled eggs 
(>6th eggs) were distributed randomly. The first instar larvae seemed to 
wander not to take eggs for nutrients, but to kill potential conspecific 
competitors. Cannibalism helped cannibals when food density was low 
(e.g., Osawa 1992), because the relatively small host R. indica is heavily 
damaged by a single larva over its complete life cycle (Yano 1993). 

Most eggs of P. rapae are deposited on the under surface of leaves 
(Yamamoto 1985 observed that 97% of females deposited eggs on the 
undersides of leaves of the field cress, R. indica). This may induce 
oviposition by more than one female on the same leaf. In fact, Watanabe 
and Yamaguchi (1993) counted 781 field cress leaves that had eggs, and 
found that 25% of them received more than one egg. This presumably 


314 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


means eggs from more than one female. However, no evidence was ob- 
tained for segregation of females on different host plants. 

Ohtani & Yamamoto (1985) found that females of P rapae have no 
site fidelity, emigrating from their emergence site. Since the females lay 
eggs singly and seldom return to deposit on the same host plant (Wata- 
nabe & Yamaguchi, unpubl. data), eggs on the same leaf may be de- 
posited by two or more females. Porter (1992) stated that single-egg-lay- 
ing females distribute eggs over a large number of host plants, and that 
this spreads the risks of predation and cannibalism, and reduces compe- 
tition with other larvae. Rothschild & Schoonhoven (1977) concluded 
that P. rapae discriminated between a cabbage leaf from which conspe- 
cific eggs had been removed and a clean control leaf. Few accounts of 
butterfly cannibalism have considered kin relatedness (e.g., Courtney 
1986), though Jones (1982) reviewed cannibalism in relation to kin se- 
lection. 


ACKNOWLEDGMENTS 


We thank K. Ueda and Y. Kato for critically reading the manuscript. The work was sup- 
ported in part by a Research Grant from the Okasan-Kato Foundation. 


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YAMAGUCHI, H. & M. WATANABE. 1993. Hatching process of pierid butterflies. II. 
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YAMAMOTO, M. 1981. Comparison of population dynamics of two pierid butterflies, Pieris 
rapae crucivora and P. napi nesis, living in the same area and feeding on the same 
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Received for publication 6 June 1996; revised and accepted 25 September 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1997, 316-332 


NOTES ON HESPERIIDAE IN NORTHERN GUATEMALA, 
WITH DESCRIPTIONS OF NEW TAXA 


GEORGE T. AUSTIN 


Nevada State Museum and Historical Society, 700 Twin Lakes Drive, 
Las Vegas, Nevada 89107, USA 


ABSTRACT. Several significant records of Hesperiidae were obtained in the vicinity 
of Parque Nacional Tikal, northern Guatemala. A new genus, Vinpeius, is proposed for 
Pompeius tinga Evans (=Vinius freemani L. Miller). Another new genus, Inglorius, is pro- 
posed for a newly described species, Inglorius mediocris. Niconiades incomptus, simi- 
lar to Niconiades xanthaphes Hiibner, is described as a new species. Range extensions are 
reported for Methionopsis dolor Evans, Mnasitheus nitra Evans, Parphorus storax (Ma- 
bille), Styriodes zeteki (Bell), Phlebodes campo Evans, Euphyes antra Evans, Amblyscirtes 
tolteca Scudder, Aides brilla (Freeman), Ridens allyni Freeman, Cyclosemia leppa Evans, 
and Staphylus lenis Steinhauser. Genitalia are illustrated for many of the foregoing, in- 
cluding variation in the harpes of Nisoniades rubescens (Méschler). 


Additional key words: Central America, distribution, genitalia, Neotropics. 


The butterflies of Guatemala are poorly known. Except for reports on 
a few old collections (Boisduval 1870, Godman & Salvin 1879-1901, 
Gibbs 1912), nearly nothing has been published on this fauna. A survey 
and monitoring study in the Parque Nacional Tikal region, Petén De- 
partment, northern Guatemala, has produced numerous interesting 
records (Austin et al. 1996), many of which represented species not pre- 
viously recorded for the country. Most of these were known from sur- 
rounding countries, but others extended distributions considerably 
southward or northward, sometimes spectacularly. Miller (1985) ob- 
served that several butterfly species exhibited apparently broad disjunc- 
tions between southern Central and South American populations and 
those in Mexico similar to those noted below. I will herein discuss sig- 
nificant extensions of known ranges among skippers (Hesperiidae), pro- 
pose two new genera of Hesperiinae, and describe two new species. 


HESPERIINAE 
Vinpeius Austin, new genus 
Type species: Pompeius tinga Evans, 1955 


Description. Palpi slender, third segment protruding about 1/2 length of second, 
pale yellow-orange with scattered black scales; antennae long, reaching beyond end of dis- 
cal cell, nearly 60% of costal length, yellow beneath club and on most of ventral surface of 
shaft except narrowly black at segments, club 1/3 shaft length, bent to apiculus after thick- 
est part, apiculus length 1.5x width of club, nudum brown, of 14 segments (6 on club, 8 
on apiculus); forewing discal cell somewhat produced anteriorly, just over 75% length of 
anal margin, vein CuAg, arises somewhat nearer origin of vein CuA, than to base of wing, 
hindwing discal cell about 1/2 width of wing; mid tibiae spined on inner surface and with 
one pair of terminal spurs, hind tibiae with two pairs of spurs; forewing somewhat pro- 
duced, costa very slightly concave just before middle, termen evenly convex, stigma along 


VOLUME 51, NUMBER 4 317 


cubitus from origin of vein CuA, nearly to origin of CuA, where bent posteriad across vein 
CuA, to 1/2 distance to vein 2A where angled proximad again nearly reaching 2A, com- 
posed of numerous fine gray hair-like scales interspersed with shorter spine-like black 
scales, these continuous along anterior edge where adjoining cubitus, entire stigma nar- 
rowly surrounded by unmodified, but semierect brown scales, these extending from 
stigma to base of cell CuA,-CuA,; hindwing evenly convex except slightly indented in cell 
CuA,-2A. Male genitalia with tegumen short, but with central spur which extends caudad 
over uncus; uncus short, blunt, broad, not divided; gnathos short, not reaching end of un- 
cus, divided, arms convergent; vinculum nearly straight; saccus moderately long; valva 
broad; costa/ampulla margin gradually ascending caudad; harpe very broad; caudal margin 
excavate ventrad, dorsal margin triangular with narrow tooth-like projection dorsad from 
inner surface barely exceeding dorsal margin; aedeagus tubular with long (about 2/5 total 
aedeagus length), narrow, and spinate caudal projection from lower right side; no comutus. 

Etymology. The name is a combination of parts of the names of the two genera in 
which the included species was previously placed, Vinius Godman, 1900 and Pompeius 
Evans, 1955. 

Diagnosis. A full diagnosis is given below under the one included species of Vin- 
peius. 


Vinpeius tinga (Evans, 1955), new combination 
(Fig. 11) 


Pompeius tinga Evans, 1955 
Vinius freemani L. Miller, 1970, new synonymy 
Pompeius freemani de la Maza et al. 1991, new synonymy 


A male hesperiine taken south of Parque Nacional Tikal and east of 
Coaba on 1 Oct. 1994 initially defied generic determination using the 
keys of Evans (1955). The genitalia of this specimen, however, resem- 
bled one species illustrated by Evans (1955), Pompeius tinga, and the 
description given in the accompanying text confirmed this identification 
(note that the Evans figures of the tegumen, uncus, gnathos, and aedea- 
gus are different from those shown herein and by Miller [1970]; the un- 
cus and gnathos of the Evans specimen were evidently lost in dissection 
and only the tegumen and aedeagus were illustrated). A further search 
of the literature from surrounding countries indicated that this taxon 
was redescribed by Miller (1970) as Vinius freemani. The Tikal male 
matches this taxon perfectly in wing pattern and genitalia. 

The characters of Vinpeius tinga are neither those of Pompeius nor 
Vinius. Evans (1955) included Pompeius in his “Hesperia Sub-group” of 
the “Hesperia Group” of hesperiines and characterized the genus as 
having the antenna nearly 1/2 the length of the costa with the club 1/4 
the length of the antennal shaft, an apiculus equalling the width of the 
antennal club, a nudum of 13 segments with 6 or 7 of these on the 
apiculus, and a well-marked black stigma on the dorsal forewing. Exam- 
ination of the type species, Pompeius pompeius (Latreille, [1824]), indi- 
cated that this diagnosis needed some embellishment. The nudum of P. 
pompeius varies from 13 to 14 segments arranged as 7 on the club and 6 


318 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(n = 8) or 7 (n = 2) on the apiculus. It should be noted here that the an- 
tennal segments vary in number and are difficult to count (e.g., Burns 
1964, MacNeill 1964), especially in distinguishing between those on the 
club and those on the apiculus (Steinhauser 1981). The blunt third seg- 
ment of the palpus barely protrudes from the scales of the second seg- 
ment. The mid tibiae are conspicuously spined on both the outer (stout, 
short) and inner surfaces (stout, long). The origin of vein CuA, is barely 
distad of the midpoint between the wing base and vein CuA,. The 
stigma is conspicuous and complex extending from the base of vein 
CuA, to nearly the base of CuA, and then posteriad to vein 2A. The an- 
terior edge along vein CuA, consists of relatively dense, small, spike-like 
black scales. Posterior to this is an area of dense, hair-like, gray scales 
which curves posteriad across CuA, and extends nearly to 2A. This gray 
area has scattered black scales and is margined posterio-distad by a nar- 
row line of spike-like black scales. A more or less round patch of these 
black scales also occurs at the posterior end of the gray area; this is the 
“lower brush patch” of MacNeill (1964). Distad of this is a large area of 
somewhat modified shiny scales extending posteriad from near the base 
of CuA,, bulging outward at CuA, and angling proximad to 2A. Similar 
scaling occurs in the base of CuA,-CuA, and in the postbasal area of 
CuA,-2A. 

The male genitalia (see figures in Godman & Salvin 1879-1901, Hay- 
ward 1951, Evans 1955) consist of a long and relatively narrow tegumen 
(V-shaped on the posterior edge in dorsal view), divided uncus with long 
and narrow arms in lateral view and narrowly pointed in dorsal view, and 
divided gnathos with long and narrow arms in lateral view with the tips 
laterad of the uncus arms in dorsal view. The vinculum is slightly curved 
and the saccus is short. The valva has a sharply sloping cephalad end, a 
prominent dorsal spike from the harpe, and the sacculus gradually nar- 
rows caudad extending nearly to the caudal end of the harpe. The 
aedeagus is tubular and the caudal end has short lower and lateral lips, 
the latter with thorn-like teeth. The two cornuti are short, tubular, and 
prominently dentate. 

Of the six additional species included by Evans (1955) in Pompeius, I 
examined three. Pompeius amblyspila (Mabille 1897) is very similar to 
P. pompeius in numerous characters including antennae (nudum 7/6), 
stigma, and genitalia (figured by Bell 1932, Hayward 1951, Evans 1955). 
Pompeius verna (Edwards 1863), including its two subspecies, P. v. 
verna and Pompeius verna sequoyah (Freeman 1942), is somewhat dif- 
ferent and may belong to another genus. The nudum is 6/7, the stigma 
is less complex and extensive and without a “lower brush patch,” and the 
genitalia (figured by Scudder 1889, Lindsey et al. 1931, Evans 1955) are 
very different, including a shorter and stouter tegumen with a shallow 


VOLUME 51, NUMBER 4 319 


V-shape cephalad in dorsal view, a shorter and blurter uncus in both lat- 
eral and dorsal view with the arms not proximate in dorsal view, the 
gnathos much narrower than the uncus, and the cornutus an inconspic- 
uous long and filament-like structure. “Pompeius” tinga is discussed 
above and below. 

The taxa included in the “Vinius Group” by Evans (1955) either lack 
androconial structures on the forewing or have brands except for one 
genus (Wahydra Steinhauser, |1991]) with a stigma. Vinius was charac- 
terized (Evans 1955) by antennae longer than 1/2 the costa length with 
the club 1/4 the length of the shaft, a nudum of 13 segments of which 
10 are on the apiculus, spined mid tibiae, males with short brands above 
and below the middle of vein CuA,, and an erectile hair tuft along vein 
3A on the dorsal hindwing with a groove in the same position of the ven- 
tral hindwing. Additional characteristics include an apiculus which is 
about 2x club width, the sharply pointed third segment of the palpus ex- 
tends beyond the scales of the second segment by about 1/4 the length 
of the second segment, the mid tibial spines are fine and on the inner 
surface, and the origin of forewing vein CuA, is much closer to CuA, 
than to the wing base. The male genitalia (e.g., figures by Godman & 
Salvin 1879-1901, Williams & Bell 1934, Evans 1955, Mielke 1968, 
Biezanko & Mielke 1973) have a short tegumen, a blunt uncus that is 
short, broad, and not divided, a short and divided gnathos with parallel 
arms, a strongly curved vinculum, and a short saccus. The valvae of 
Vinius are variable with the harpe caudally sloping or having a toothed 
dorsal margin. The aedeagus is tubular and with no prominent caudal 
extensions or cornutus. 

The antennae of Vinpeius tinga are proportionately longer than on P. 
pompeius and about the same as on Vinius and the antennal club is 
longer than on either genus. The nudum of Vinpeius has 14 segments 
(one more than either Pompeius or Vinius) with eight of these on the 
apiculus (more than the 6 or 7 on Pompeius and less than the 10 on 
Vinius). The third segment of the palpus protrudes from the second 
much more than on either Vinius or Pompeius. Vinpeius has fine mid 
tibial spines only on the inner surface as on Vinius and not stout spines 
on both the inner and outer surfaces as on Pompeius. The forewing dis- 
cal cell of Vinpeius is shorter in relation to the anal margin than on ei- 
ther Pompeius or Vinius (over 80% its length on both genera). The ori- 
gin of forewing vein CuA, is intermediate between the origin of this vein 
on Pompeius and Vinius. The androconial structure of Vinpeius extends 
across wing cells and thus is a stigma rather than brands which are sim- 
pler structures and parallel to veins. The species of Vinius obviously 
have brands, these in an unusual position over and under the middle of 
vein CuA, as noted by Evans (1955). The structure of the stigma on Vin- 


320 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


peius is very different from those on the Pompeius species examined, 
being much less complex. Vinpeius lacks the prominent hindwing hair 
tuft present on Vinius. The genitalia of Vinpeius differ in several re- 
spects from both Vinius and Pompeius, especially in the form of the 
aedeagus. 

No other genus has the combination of characters seen on Vinpeius; 
it will not key to any of the eight Hesperiine group keys in Evans (1955) 
despite his inclusion of V. tinga in Pompeius. In the “Hesperia Group” 
key V. tinga will key to the “Phemiades Sub-group” with “Nudum of 14 
or more segments.” Even if it remained in the “Hesperia Sub-group” in- 
cluding Pompeius, there are not more nudum segments on the club than 
on the apiculus. Vinpeius has too many segments to the nudum to key 
to any “Vinius Group” genus in Evans (1955). 

The relationships of Vinpeius are, at best, unclear. Its stigma some- 
what suggests that among the genera of the last half of Evans’ (1955) 
“Hesperia Group” taxa, its antennal structure suggests the “Vinius 
Group” or “Apaustus Group”, and the general color and pattern is char- 
acteristic of both the “Vinius” and “Hesperia” groups. The problems 
with Evans’ (1955) often artificial groupings have been reiterated (e.g., 
Burns 1990) and the previous inclusion of V. tinga in both Vinius and 
Pompeius further demonstrates these problems. For the present, place- 
ment of Vinpeius among the “Vinius Group” taxa should suffice. | 


Inglorius Austin, new genus 
Type species: Inglorius mediocris Austin, new species 


Description. Palpi slender, third segment straight, protruding well beyond second 
segment, about equal to length of dorsal edge of second segment; antennae long, extend- 
ing beyond end of forewing discal cell, nearly 60% length of forewing costa, black with 
pale ochreous beneath distad and below club; club just over 1/4 (28%) antennal length, 
bent to apiculus at thickest part, apiculus length about 2x club width, nudum gray, of 12 
segments (3 on club, 9 on apiculus); forewing discal cell slightly produced, 75% length of 
anal margin, origin of vein CuA, nearer to CuA, than to wing base, hindwing discal cell 
just over 1/2 wing width; mid tibiae with four fine spines on inner surface and single pair 
of spurs, hind tibiae with two pairs of spurs; forewing produced with slight concavity be- 
tween CuA, and 2A; hindwing convex anteriorly, somewhat concave between CuA, and 
2A; no apparent secondary sexual characters. Male genitalia with short tegumen; uncus 
longer than tegumen, undivided, and hoodlike over gnathos; gnathos as long as uncus, di- 
vided, extending laterad of uncus in dorsal view and as rectangular flaps mesad in ventral 
view; vinculum sinuate; saccus short; valva very long, ampulla/costa long and sloping some- 
what downward caudad, harpe long, roughly triangular ending in an inward turned point 
caudad, dorsal margin undulate, weakly serrate cephalad; aedeagus tubular (anterior por- 
tion missing), caudal end expanded terminally in lateral view, no apparent cornutus. 

Etymology. The name means “undistinguished,” as the only known species of the 
genus is a nondescript brown insect. 

Diagnosis. Inglorius appears to belong within Evans’ (1955) “Apaustus Sub-group” 
of his “Apaustus Group” characterized by a long third segment of the palpi. Most of these 
fourteen genera contain brown species with few distinguishing marks. None of these, nor 
any other hesperiine, has the combination of characters seen on Inglorius as outlined 


VOLUME 51, NUMBER 4 321 


Fics. 1-6. Hesperiidae from northern Guatemala (dorsum on left, venter on right; all 
from GUATEMALA: Petén; Parque Nacional Tikal, unless noted otherwise). 1, Euphyes 
antra, male (25 June 1993). 2, E. antra, female (30 July 1992). 3, Styriodes zeteki, male 
(15 July 1993). 4, Inglorius mediocris, holotype male. 5, Aides brilla, female (29 Dec. 
1992). 6, Staphylus lenis, female (south of Parque Nacional Tikal, east of Cauba, 1 Oct. 
1994). 


above. The genitalia are particularly unique and totally unlike those of any other known 
taxon. 


Inglorius mediocris Austin, new species 
(Figs. 4, 12) 


Description. Male: forewing length of holotype = 11.8 mm; in addition to generic 
description above, dorsum brown, scattered ochreous scales, these forming vague macules 


322 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 7-10. Niconiades species (dorsum on left, venter on right). 7, N. incomptus, 
holotype male. 8, N. xanthaphes, male (BRAZIL: Rondénia; 62 km S of Ariquemes, linha 
C-20, Fazenda Rancho Grande, 27 Nov. 1991)..9, N. incomptus, paratype female 
(GUATEMALA: Petén; El Remate, Cerro Cahui, 29 Sept. 1994). 10, N. xanthaphes, fe- 
male (same location as Fig. 8, 14 Nov. 1992). 


(not seen except under magnification) in CuA,-CuA, just beyond origin of CuA,, smaller 
macules offset distad in M3-CuA, and in upper portion of discal cell; long ochreous hair- 
like scales on forewing at base of CuA,-2A and along basal 1/2 of anal margin; hindwing 
immaculate with ochreous hairlike scales on posterior 1/2; fringes of both wings very worn, 
appearing gray. Ventral forewing paler brown especially distad, slight purplish cast along 
costa; hindwing with similar purplish cast over most of wing except for brown anal fold, 
small cream-colored macules at distal end of discal cell and as postmedian row from Rs to 
CuA,. Head brown with scattered ochreous scales especially around eyes; palpi gray with 
scattered white scales beneath becoming white and then ochreous on sides; thorax brown 
with scattered ochreous scales above, whitish beneath, legs pale brown; dorsal abdomen 
brown, ventral abdomen white (possibly with dark central line). Genitalia: see generic de- 
scription above. Female: unknown. 

Type. Holotype ¢ with the following labels: white, printed - Tikal, peten / Guatemala / 
September 12, 1993 / D. L. Lindsley; printed and handprinted - Genitalia Vial / GTA - 
5283; red, printed - HOLOTYPE / Inglorius mediocris / Austin. The holotype will be de- 
posited in the Entomological Collections at the Universidad del Valle, Guatemala City, 
Guatemala. Type locality. GUATEMALA: Petén; Parque Nacional Tikal. 

Etymology. The name means “ordinary” as this is a rather ordinary brown skipper. 


VOLUME 51, NUMBER 4 323 


14 


Fics. 11-14. Genitalia of male Hesperiidae; all from GUATEMALA: Petén. 11, Vin- 
peius tinga, GTA Vial #5230 (lateral view of uncus, gnathos, tegumen, vinculum, saccus; 
internal view of right valva; right and left lateral and dorsal views of aedeagus; dorsal and 
ventral views of uncus, gnathos, and caudal end of tegumen). 12, Inglorius mediocris, 
holotype, GTA Vial #5283 (lateral view of uncus, gnathos, tegumen, vinculum, saccus; in- 
ternal view of right valva; dorsal and left views of caudal end of aedeagus; dorsal view of 
uncus, gnathos, and tegumen; ventral view of uncus and gnathos). 13, Mnasitheus nitra, 
GTA Vial #3236 (lateral view of uncus, gnathos, tegumen, vinculum, saccus; internal view 
of right valva; left and dorsal views of aedeagus; dorsal and ventral views of uncus, gnathos, 
and tegumen; ventral view of juxta). 14, Euphyes antra, GTA Vial #5190 (lateral view of 
uncus, gnathos, tegumen, vinculum, saccus; internal view of right valva; left and dorsal 
views of aedeagus; dorsal and ventral views of uncus, gnathos, and tegumen). 


324 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Diagnosis and discussion. The type of Inglorius mediocris, a worm male of a small, 
nearly entirely brown, skipper does not resemble any described genus or species (see also 
generic diagnosis above). The species is known only from the holotype taken in mid Sep- 
tember. 


Methionopsis dolor Evans, 1955 


This species has previously been reported from Panama southward 
(Evans 1955). Single males were taken at Tikal by D. Lindsley on 11 and 
12 Sept. 1993. 


Mnasitheus nitra Evans, 1955 
(Fig. 13) 


Evans (1955) described M. nitra based on a pair of males from 
Parana, Castro (southern Brazil). There seems to be no subsequent re- 
port of this species although it is known from Peru (fide O. Mielke). It 
thus was a surprise to find it among the Tikal area fauna. Three males, 
all from Parque Nacional Tikal, were taken on: 29 Feb. 1992, leg. N. M. 
Haddad; 31 May 1992, leg. N. M. Haddad, and 8 June 1994, leg. G. A. 
Orellana. The genitalia of one of these is illustrated. 


Parphorus storax storax (Mabille, 1891) 


Evans (1955) recorded the distribution of this species as from Costa 
Rica southward. Monroe and Miller (1967) reported a record for Hon- 
duras. It was not known from EI! Salvador (Steinhauser 1975) nor Mex- 
ico (de la Maza et al. 1991). A male from Parque Nacional Tikal taken 
on 4 Feb. 1992 by G. T. Austin represents a northward extension of its 
known distribution. 


Styriodes zeteki (Bell, 1931) 
(Figs. 3, 15) 


This species was described from a single male taken on Barro Colorado 
Island in the Canal Zone, Panama (Bell 1931), not from Bolivia as stated 
by Evans (1955) and has not otherwise been reported. Two males from 
Tikal taken on 15 July 1993, leg. G. A. Orellana and 12 Sept. 1993, leg. D. 
Lindsley represent a considerable extension of the known distribution. 
The male genitalia are illustrated here in more detail than previously. 


Phlebodes campo sifax Evans, 1955 


This species has only been known from South America and as far 
north as Guyana (Evans 1955). A male from Parque Nacional Tikal 
taken on 31 May 1992 by N. M. Haddad represents a significant exten- 
sion of the reported distribution. 


VOLUME 51, NUMBER 4 325 


Fics 15-19. Genitalia of male Hesperiidae; all from GUATEMALA: Petén, unless 
noted. 15, Styriodes zeteki, GTA Vial #4699 (same structures as Fig. 14). 16, Cyclosemia 
leppa, GTA Vial #1992 (same structures as Fig. 13). 17, Niconiades incomptus, holotype, 
GTA Vial #5171 (same structures as Fig. 13). 18, Niconiades xanthaphes, GTA Vial #2524 
from same location as Fig. 8 (lateral view of uncus, gnathos, tegumen, vinculum, saccus; 
internal view of right valva; left and dorsal views of aedeagus). 19A, Nisoniades rubescens, 
GTA Vial #5147 (internal view right and left valvae; flattened view of caudal end of right 
harpe). 19B, Nisoniades rubescens, GTA Vial #5135 (same structures as Fig. 19A). 


Euphyes antra Evans, 1955 
(Figs. 1, 2, 14, 20) 


This species was described based on one male from Lima, Peru, and 
two putative females from “Lower Amazons” (Evans 1955). Mielke 
(1972) found that the two females were of another taxon, Euphyes de- 
rasa tuba Evans, 1955, and knew of no other records of E. antra. An in- 
dication of how little we know of Neotropical hesperiid faunas was the 
discovery of a male and female of E. antra among the Tikal material 


326 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


taken in the park on 25 Jan. 1993 and 30 July 1992, respectively, by G. 
A. Orellana. Since Mielke (1972) was unable to illustrate all the struc- 
tures of the male genitalia, these are fully illustrated here. The female 
has not been previously described. It is similar to the male with broader 
and less produced wings and the pale yellow median band on the ven- 
tral hindwing is broader. The genitalia are most similar to those of 
Mielke’s (1972) “subferruginea” and “peneia” groups with, especially, 
the long and relatively thin ductus bursae. 


Amblyscirtes tolteca tolteca Scudder, 1872 


Freeman (1993) recently reviewed the distribution of this species and 
indicated a distribution over most of Mexico including Chiapas. Stein- 
hauser (1975) tentatively included this species for E] Salvador but the 
record has not been verified. A single male from Parque Nacional Tikal 
taken on 14 July 1992 and two more taken on 18 July 1992 by G. A. 
Orellana represent a new record for Guatemala and an eastern exten- 
sion of the species’ distribution. 


Niconiades incomptus Austin, new species 
(tes, 7, ©) la, Zl) 


Description. Male: forewing length = 17.1 mm (holotype), 17.0 mm (paratype); 
forewing produced, apex moderately rounded, termen slightly concave in CuA,-2A; hind- 
wing narrow, apex rounded, termen concave to prominently elongate tornus; dorsum 
blackish brown with prominent blue-green in basal 1/3 of CuA,-2A and basal 1/2 of anal 
margin of forewing and on basal 1/2 of hindwing; forewing with three short gray brands, 
one above the other, one above vein CuA, at base of cell (the broadest), one below CuAg, 
and another above 2A; forewing with very pale yellow hyaline macules as follows: in discal 
cell, strongly constricted in middle nearly separating upper and lower portions; CuA,-2A, 
semicircular in lower half of cell over middle of vein 2A; mid CuA,-CuAg, square with 
slightly excavate distal edge; M3-CuA,, more or less quadrate, smaller than and offset dis- 
tad from or contiguous with that in CuA,-CuA,; subapical, aligned in R3-Ry, Ry-Rs, Rs-M;, 
rectangular, that in R,-R, smallest; fringe dark gray anteriorly, white behind vein CuAg; 
hindwing with very pale yellow, more or less rectangular hyaline macules in M3-CuA, and 
CuA,-CuAg,; fringe brown at apex and tornus, otherwise white. Venter blackish brown, 
paler brown distad; forewing with macules repeated from dorsum, that in CuA,-2A more 
quadrate, extended, especially distad, by white scaling, elongate cream-colored macule an- 
terior to discal cell macule in Sc-R, and R,-Rg, this extending basad as sparse scaling to 
wing base in Sc-R,, similar scaling in base of costal cell; hindwing with hyaline macules 
outlined with white; narrow white band with ill-defined margins from costa (where vague) 
posteriad (including hyaline macules) to vein 2A where broadest and hooked somewhat 
basad. Dorsum of head, thorax, and anterior abdomen blue-green, posterior abdomen 


=> 


Fics. 20-25. Genitalia (ventral view, including lamellae, antrum, ductus bursae, cor- 
pus bursae) of female Hesperiidae (all from GUATEMALA: Petén, unless noted). 20, Eu- 
phyes antra, GTA Vial #6318. 21, Niconiades incomptus, paratype, GTA Vial #5172. 22, 
Niconiades xanthaphes, GTA Vial #3128, from BRAZIL: Rondénia. 23, Ridens allyni, 
GTA Vial #5281. 24, Staphylus lenis, GTA Vial #5298, #5290, #5300, #5301 (lamellae). 25, 
Aides brilla, GTA Vial #5282 (including papillae anales). 


327 


VOLUME 51, NUMBER 4 


328 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


dark brown; palpi blue-green above, whitish beneath and on cheeks; antenna black, 
vaguely white at base of club beneath, nudum dark gray with 17, 18 segments; pectus 
bright ochreous with green tinge; legs dark brown with some pale ochreous scaling, mid 
tibiae spined and with single pair of spurs, hind tibiae with two pairs of spurs; ventral ab- 
domen whitish with very broad black median band. Genitalia: tegumen bulbous; uncus un- 
divided, broad, blunt; gnathos slightly shorter than uncus, divided, pointed caudad, arms 
somewhat convergent; vinculum slightly sinuate; saccus short, upturned; valva broad, 
harpe stout, triangular, dorso-caudal margin with irregular fine serrations, produced to 
short tooth just after ampulla, ventral margin narrowly excavate, prominent “shelf” pro- 
truding mesad from upper inner surface; juxta as narrow band, ventral portion trifurcate 
cephalad, large cluster of bristles dorsad; aedeagus tubular, narrowing caudad to lateral 
prongs ventrad, left prong slightly longer than right; no comutus. Female: forewing length 
17.0, 18.8, 19.8 mm; similar to male; no brands; forewing discal cell macule less con- 
stricted in middle; forewing broader; hindwing termen less concave; antennal nudum with 
18, 18, 19 segments. Genitalia: lamella postvaginalis not well developed; lamella antevagi- 
nalis with long central process extended caudad where bifurcate; ductus bursae and cor- 
pus bursae not clearly separable, gradually expanding to bulbous cephalad end. 

Types. Holotype 4 with the following labels: white, printed - GUATEMALA / Petén, 
E] Remate / Cerro Cahui / 30 Sept. 1994 / leg. G. T. Austin; white, printed and hand- 
printed - Genitalia Vial / GTA - 5171; red, printed - HOLOTYPE / Niconiades incomptus 
/ Austin. Paratypes - same location as holotype, 29 Sept. 1994, leg. G. A. Orellana (1 3); 28 
Sept. 1994, leg. G. T. Austin (1 @); 29 Sept. 1994, leg. G. T. Austin (1 2); Parque Nacional 
Tikal, 25 Mar. 1992, leg. N. M. Haddad (1 2); 26 Sept. 1992, leg. J. V. Orellana (1 2). The 
holotype and a female paratype will be deposited in the Entomological Collections at the 
Universidad del Valle, Guatemala City, Guatemala. Type locality. GUATEMALA: Petén; 
E] Remate, Cerro Cahui. This is on the north shore of Lago Petén Itza with a mosiac of 
mature and second growth forests. Most of the types were taken along the forest edge. 

Etymology. The name means “untrimmed” referring to the ventral hindwing white 
band without well-defined edges. 

Distribution. The distribution of this species is currently unknown and some of the 
Niconiades xanthaphes Hiibner, [1821] reported from Central America and elsewhere 
may refer to N. incomptus. A pair of N. incomptus was seen from the Atlantic Slope of 
Costa Rica (male from Limon Province, female from Heredia Province); another pair was 
seen from the vicinity of Candelaria, Oaxaca, Mexico. All reports of N. xanthaphes from 
Mexico south through Central America should be treated as suspect until the specimens 
are reexamined. Certainly, N. incomptus is more widespread than the records indicated 
above and is residing in collections among series of N. xanthaphes. 

Diagnosis and discussion. This new species is most similar to N. xanthaphes which 
may be slightly smaller in size (male forewing length = 16.7 mm [15.9-17.9, n = 10], fe- 
male forewing length = 16.6, 17.3, samples from Rond@6nia, Brazil). The forewing of N. 
xanthaphes (Figs. 8, 10) is stouter and less produced apically than on N. incomptus (Figs. 
7, 9), the hindwing is broader and less concave with a shorter anal lobe, the ventral 
forewing has the macules anterior to the discal cell macule more distinct and pale yellow- 
orange, the hindwing band is broader, nearly the full width of the hyaline macules, and 
with margins sharply defined, and there is a narrow white streak along the distal 1/3 to 1/2 
of vein 3A. 

While N. incomptus is readily separable from N. xanthaphes by characters of the wings 
(Figs. 7-10), the genitalia of the two species are very similar. The male genitalia of N. in- 
comptus do not appear separable from those of N. xanthaphes (Fig. 18); those illustrated 
by Godman and Salvin (1879-1901), Hayward (1951), and Evans (1955) could be of ei- 
ther species. The female genitalia are also very similar but the central process of the 
lamella antevaginalis on N. incomptus is less robust than on N. xanthaphes (Fig. 22). The 
forewing with brands illustrated by Godman and Salvin (1879-1901) appears to be N. xan- 
thaphes based upon the more rounded and less produced apex; they may have seen both 
species as there is no mention of the white streak on the anal margin of the ventral hind- 
wing. Hayward (1951) mentioned this character and undoubtedly saw N. xanthaphes from 
Argentina. Mielke (pers. comm.), in review of this manuscript, suggested that N. incomp- 


VOLUME 51, NUMBER 4 329 


tus was a northern subspecies of N. xanthaphes, but he indicated overlap between the two 
in Panama. 


Aides brilla (Freeman, 1970) 
(Figs. 5, 25) 


A male from Tikal (16 Sept. 1993, leg N. M. Haddad) is like the sin- 
gle previously reported specimen, the holotype male fom Catemaco, Ve- 
racruz, Mexico (Freeman 1970). An additional Aides from Tikal (29 
Dec. 1992, leg. G. A. Orellana) is apparently the first known female A. 
brilla. The wings are more elongate than on the male (forewing length 
= 26.5 mm) and the dorsal color and pattern is virtually identical except 
the discal cell macule is further from the macule in CuA,-CuA,. The 
ventral forewing is similar to that of the male as is the color of the ven- 
tral hindwing. The silverly-white maculation of the ventral hindwing, 
however, differs. The large macule in CuA,-2A is similar in shape but 
does not extend as far basad, there is no macule in the base of CuA,- 
CuA,, the discal cell macule is a small round spot, similar (slightly 
larger) spots occur in the middle of M,-M, and submargin of M,-CuA,, 
and an additional oval macule is in the submargin of CuA,-CuA,. The 
genitalia of this female are illustrated. 


PYRGINAE 


Ridens allyni Freeman, 1979 
(Fig. 23) 


This species is known from Veracruz, Oaxaca, and Chiapas, Mexico 
(Freeman 1979). It is not uncommon in the Tikal region with records for 
11 Mar. 1993, leg. G. A. Orellana (1 male), 13 July 1992, leg. G. A. Orel- 
lana (1 female), 18 July 1992, leg G. A. Orellana (1 male), 23 Aug. 1993, 
leg. J. V. Orellana (1 female), and 25 Sept. 1992, leg. G. A. Orellana (1 
male). The female genitalia are illustrated for the first time herein. 


Nisoniades rubescens (Moschler, [1877]) 
(Fig. 19) 


Eight male Nisoniades Hiibner, 1819 from Tikal were identified as N. 
rubescens with the key in Evans (1953). These, however, exhibit two 
somewhat different configurations of the valvae, especially the right. 
One phenotype, represented by a single specimen (Fig. 19A), is that il- 
lustrated as N. rubescens by Evans (1953) or its putative synonym Pelli- 
cia bromias Godman & Salvin, [1894] illustrated by Godman & Salvin 
(1879-1901) and Hayward (1948). On this, the ampulla/costa of the 
right valva is broadly and evenly convex, the caudal end of the ampulla 
has a somewhat upward orientation, and the harpe is broadly rounded 


caudad. The harpe of the left valva is rather sharply bent. The remaining 


330 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


seven individuals are of the second phenotype (Fig. 19B) and has not 
been previously illustrated. The costa/ampulla of the right valva is less 
evenly convex, the caudal end of the ampulla has more of a ventrad orien- 
tation, and the caudal end of the harpe is truncated to a long and narrow 
finger-like lobe. The harpe of the left valva is less sharply bent. 

No superficial differences between the two could be detected. Both 
genitalic phenotypes have also been seen among specimens from Costa 
Rica. Examination of additional material (including females) and of the 
types of N. rubescens and its listed synonyms P. bromias, Pellicia clara 
Mabille & Boullet, [1917], Pellicia nigra Mabille & Boullet, [1917], and 
Achlyodes triangulus Mabille, 1897 are required to properly evaluate 
the observed variation; two species may be involved. Evans (1953) noted 
variation in the genitalia of Nisoniades maura (Mabille & Boullet, 
[1917]), Nisoniades mimas (Cramer, [1775]), and Nisoniades ephora 
(Herrich-Schiffer, 1870). 


Cyclosemia leppa Evans, 1953 
(Fig. 16) 


This species is evidently known only from the holotype male from Bo- 
livia and a female from Peru (Evans 1953). A single very worn male 
taken at Tikal on 4 Feb. 1992 by G. T. Austin represents a major range 
extension. Its genitalia are illustrated herein. 


Staphylus lenis Steinhauser, 1989 
(Figs. 6, 24) 


This species is relatively common in the Tikal region with records for 
February and May through October. At the time of its description, S. le- 
nis was known only from males taken in Quintana Roo, Mexico and in 
Trinidad (Steinhauser 1989). The female (forewing length = 12.4 mm 
[11.7-12.8 mm, N = 4]) is similar to females of other species of the 
Staphylus mazans (Reakirt, [1867]) group, especially Staphylus ascala- 
phus (Staudinger, 1875) and Staphylus unicornis Steinhauser and 
Austin, 1993. It differs from female S. ascalaphus (forewing length from 
Costa Rica = 12.9 mm [12.2-13.3 mm, N = 10]) by its smaller mean size 
and less prominent contrast between the brown ground color and the 
blackish bands on the dorsum. It differs from the slightly larger female 
S. unicornis (forewing length from Costa Rica = 12.7 mm [12.0—13.9 
mm, N = 12]) by the absence of the lower hyaline macule in the 
forewing discal cell (this present on most S. unicornis). Two of four fe- 
males of S. lenis lack a white macule in M,-CuA,; this is absent on most 
S. ascalaphus but present on nearly all S. unicornis. 

The lamellae of the female genitalia of S. lenis are highly variable 
(Fig. 24) as also shown for S. ascalaphus and S. unicornis by Steinhauser 


VOLUME 51, NUMBER 4 331 


(1989) and Steinhauser and Austin (1993). Generally, the plate-like 
lamella postvaginalis of S. lenis has a roughly heart-shaped and heavily 
sclerotized central process (this varies in size and shape) on its caudal 
edge, flanked by usually oval membranous lobes (also variable in size) 
with microtrichia especially caudad. The lamella antevaginalis has a cau- 
dally excavate central plate with two large and caudally pointed lateral 
processes and central serrations; the depth of the central concavity and 
number of serrations varies. No other S. mazans group species exam- 
ined have the prominent lateral processes of the lamella antevaginalis 
seen on S. lenis (Steinhauser 1989, Steinhauser & Austin 1993). 


ACKNOWLEDGMENTS 


Nick M. Haddad, Claudio Méndez, Thomas D. Sisk, Dennis D. Murphy, Alan E. 
Launer, and Victor and Gustavo Orellana contributed greatly with their assistance in the 
field. D. L. Lindsley allowed us to examine critical specimens in his collection. Stephen R. 
Steinhauser assisted in the identification of several skippers and critically read the manu- 
script. I thank Olaf H. H. Mielke for sharing his extensive knowledge of Neotropical Hes- 
periidae. Thanks also are extended to Jack Schuster and Inio Cano of the Universidad del 
Valle for laboratory facilities and providing work and storage space in Guatemala City and 
to N. M. Haddad for comments on the manuscript. Three reviewers, J. M. Burns, C. D. 
MacNeill, and O. H. H. Mielke, made several useful suggestions for improvement of the 
manuscript. Many institutions have graciously supported efforts to understand the biology 
and conservation of butterflies in Guatemala and issued the necessary permits: Centro de 
Estudios Conservacionistas (CECON) of the Universidad de San Carlos, the Parque Na- 
cional de Tikal, The Peregrine Fund, and Consejo Nacional de Areas Protegidas 
(CONAP). Financial support for field studies was provide by the John D. and Catherine T. 
MacArthur Foundation through Stanford University. 


LITERATURE CITED 


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Park area of Guatemala. Trop. Lepid. 7:21—37. 

BELL, E. L. 1931. A list of Hesperiidae from Barro Colorado Island, Canal Zone, and ad- 
jacent Panama, with description of a new species. J. New York Entomol. Soc. 
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. 1932. Hesperiidae (Lepidoptera, Rhopalocera) of the Roraima and Duida Expe- 
ditions, with descriptions of new species. Amer. Mus. Novit. 555:1—16. 

BIENZANKO, C. M. & O. H. H. MIELKE. 1973. Contribuicao ao estudo faunistico dos 
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ondmicas e descrigGes de espécies novas (Lepidoptera). Acta Biol. Par., Curitiba 
2-102. 

BOISDUVAL, J. B. A. D. DE. 1870. Considerations sur des lépidoptéros envoyés du 
Guatemala a M. de l’Orza. Rennes, Oberthiir et fils. 100 pp. 

BURNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. 
Publ. Entomol. 37:1—214. 

. 1990. Amblyscirtes: problems with species, species groups, the limits of the 
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tion of Evans (Hesperiidae). J. Lepid. Soc. 44:11—27. 

DE LA MAZA E., J., A. WHITE L. & R. DE LA MAZa E. 1991. La fauna de mariposas de 
Mexico. Parte II. Hesperioidea (Lepidoptera: Rhopalocera). Rev. Soc. Mex. Lepid. 
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EVANS, W. H. 1953. A catalogue of the American Hesperiidae in the British Museum 


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(Natural History). Part III (groups E, F, G) Pyrginae. Section 2. London: British Mu- 

seum (Natural History). 246 pp. 

. 1955. A catalogue of the American Hesperiidae in the British Museum (Natural 
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idoptera). J. New York Entomol. Soc. 78:88—99. 

. 1979. Nine new species and seven new records of Mexican Hesperiidae. Bull. 

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. 1993. Notes on Amblyscirtes Scudder, with the description of two new sub- 
species (Insecta: Lepidoptera: Hesperiidae: Hesperiinae). Ann. Carnegie Mus. 
62:341—350. 

GipBs, A. E. 1912. Butterflies from British Honduras and Guatemala. Proc. Entomol. 
Soc. London 1912(2):xiv—xlviii. 

GopMaAN, F. D. & O. SALVIN. 1879-1901. Biologia Centrali-Americana. Zoologia, Lepi- 
doptera-Rhopalocera. London. 3 vols. 

HAYWARD, K. J. 1948. Genera et species animalium argentinorum. Vol. 1: Insecta, Lepi- 
doptera, Pyrrhopyginae et Pyrginae. Buenos Aires, G. Kraft Ltd. 389 pp. 

1951. Genera et species animalium argentinorum. Vol. 2: Insecta, Lepidoptera, 
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ica. J. Sci. Lab., Denison Univ. 26:1—142. 

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brasileiros I. Resultados de uma excursao 4 Foz do Iguacu, Paranda, Brasil, com notas 
taxonOmicas (Lepidoptera). Atas Soc. Biol. Rio de Janeiro 12:73—78. 

. 1972. As espécies Sul-Americanas do género Euphyes Scudder, 1872 (Lepi- 
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the description of a new subspecies from Mexico. Insecta Mundi 1:43—45. 

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. 1981. A revision of the proteus group of the genus Urbanus Hubner. Lepi- 

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Am. Entomol. Soc. 60:265—280. 


Received for publication 20 May 1996; revised and accepted 10 September 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1997, 333-343 


LIFE HISTORY AND BEHAVIOR OF SYNAXIS CERVINARIA 
(GEOMETRIDAE), A DEFOLIATOR OF ARCTOSTAPHYLOS 
PATULA (ERICACEAE) 


MICHAEL A. VALENTI!, ALAN A. BERRYMAN 


Department of Entomology, Washington State University, 
Pullman, Washington 99164, USA 


AND 


GEORGE T. FERRELL 


USDA Forest Service, Pacific Southwest Forest & Range Research Station, 
2400 Washington Avenue, Redding, California 96001, USA 


ABSTRACT. The life stages and behavior of Synaxis cervinaria are described from 
laboratory and field studies conducted in Shasta Co., California, using greenleaf man- 
zanita, Arctostaphylos patula, as a host plant. Instars 1 and 2 resemble the green surface 
and red edges of the expanding leaves, and third instars begin to resemble twigs. The first 
three instars remain in the crown during the day. Instars 4 and 5 resemble twigs and 
stems, maintaining stick-like resting positions near the main stem below the crown during 
the day. The pupae overwinter. Adults emerge in late spring/early summer and oviposition 
occurs prior to and during manzanita leaf expansion. There is one generation per year. 


Additional key words: greenleaf manzanita, crypsis, flight period, leaf consumption. 


Synaxis cervinaria (Packard) (Geometridae) was described from an 
adult male collected in West Springs, California (Packard 1871). 
McGuffin (1987) described the adult female and genitalia of both sexes 
but noted that the egg, early larval instars, and pupa were unknown: a 
description based on a single preserved specimen of a “mature” larva 
(presumably fifth instar) was given. Synaxis cervinaria ranges from 
British Columbia south to California, and the reported hosts include: 
Oregon white oak, Quercus garryana Hooker (Fagaceae) (Jones 1951); 
an unidentified Quercus; poplar, Populus sp. (Populaceae) (McFarland 
1965); Arbutus (Ericaceae) (McGuffin 1987); willow (Salicaceae); bitter- 
brush (Rosaceae); cascara and species of Ceanothus (Rhamnaceae) 
(Miller 1995). 

In 1990, as part of a broader study of the insect fauna associated with 
Arctostaphylos patula E. Greene (Ericaceae) (Valenti 1994), we found 
larvae of S. cervinaria on this plant near Hat Creek, Shasta Co., Califor- 
nia. This manzanita commonly occurs throughout the western United 
States in montane forest zones (Ball et al. 1983) and is of concern to for- 
est land managers because it often inhibits the survival, regeneration, 
and growth of conifers (Radosevich 1984). In response to a general lack 
of information regarding insect fauna associated with chaparral commu- 


‘Current address: Delaware Department of Agriculture, 2320 South DuPont Highway, 
Dover, Delaware 19901, USA 


334 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


nities (Force 1990), especially species of manzanita (Haws et al. 1988), 
we conducted a study of S. cervinaria and its interactions with greenleaf 
manzanita. 

This paper presents life history and behavioral information for S. cer- 
vinaria, one of six geometrids encountered during our faunal survey 
(Valenti & Zack 1995) that utilizes greenleaf manzanita as a host plant. 
The goals of this study were to: 1) determine adult flight period; 2) de- 
scribe larval stages and behavior and determine stadium lengths; 3) 
measure leaf consumption of A. patula by larvae; 4) establish a list of ac- 
ceptable foodplants associated with greenleaf manzanita communities in 
the Hat Creek area; and 5) identify natural enemies of eggs and larvae. 


MATERIALS AND METHODS 


Synaxis cervinaria laboratory studies, rearings, and adult collections 
were conducted at the Forest Insect Laboratory, USDA Forest Service, 
Hat Creek, California. Field studies were conducted at two sites in 
Shasta Co., California. The first site was adjacent to the USDA Forest 
Service Work Center at Hat Creek (T34N R4E S16) (elev. 1018 m) and 
contained a variety of woody plant species. Greenleaf manzanita ac- 
counted for less than 30% (canopy coverage) of the total vegetation. 
Other common plants included: sagebrush (Artemesia tridentata Nut- 
tall) (Compositae), curl-leaf mountain-mahogany (Cercocarpus ledi- 
folius Nuttall), birch-leaf mountain-mahogany (C. betuloides Torrey & 
A. Gray), antelope bitterbrush (Purshia tridentata [Pursh| de Candolle) 
(Rosaceae), California black oak (Quercus kelloggii Newberry) (Fa- 
gaceae), ponderosa pine (Pinus ponderosa Lawson), sugar pine (P. lam- 
bertiana Douglas), white fir (Abies concolor [Gordon & Glendinning | 
Lindley), Douglas-fir (Pseudotsuga menziesii [Mirbel] Franco), incense 
cedar (Calocedrus decurrens [Torrey] Florin), and western juniper (Ju- 
niperus occidentalis Hooker) (Pinaceae). The second site (elev. 1512 m) 
was 3.2 km northwest of California Route 89 near Logan Lake in Old 
Station, ca. 24 km south of Hat Creek (T32N R4E S2&3). Dominant 
vegetation (canopy coverage) consisted of greenleaf manzanita (74%), 
tobacco brush (11%), Ceanothus velutinus Hooker (Rhamnaceae), and 
ponderosa pine (9%). This 50+ ha site is on an east-facing slope and was 
mechanically cleared of all vegetation in 1976 and planted with pon- 
derosa pine in 1977. 

During 1991 and 1992, a 15 watt black light was used at the Hat 
Creek Forest Insect Laboratory to determine the flight period of S. 
cervinaria. All adults collected from 2100—2300 h each evening were 
tallied, a portion of the males was retained as vouchers, and most fe- 
males were placed in covered 236 ml plastic cups for egg collection. Be- 


VOLUME 51, NUMBER 4 335 


havioral data, including egg-laying, were obtained by observing adults in 
the field and in screen cages (45 x 45 x 75 cm). 

Larval rearing containers were constructed from clear plastic cylindri- 
cal tennis ball containers 20 cm in height and 7 cm in diameter. A single 
neonate larva was placed on a greenleaf manzanita branch in each of 30 
containers. Larval development under ambient conditions was observed 
daily until pupation. Foliage was replaced once, following the fourth in- 
star molt. Frass of the first four instars was collected from the bottom of 
each rearing container when foliage was changed, and again after pupa- 
tion. All leaves damaged by the fifth instars were collected, pressed, 
mounted to sheets of standard card stock, and missing portions of each 
leaf were drawn freehand. A digitizer was then used to measure leaf area 
consumed (cm?). Frass weight was used to estimate leaf area consump- 
tion for first through fourth instars. Head capsule widths and total body 
lengths of 30 individuals in each instar (reared in 45 x 45 x 75 cm cages) 
were measured to the nearest 0.05 mm using an ocular micrometer. 

A host suitability study was conducted by placing 15—40 unfed first in- 
stars onto foliage from various plants in the families Pinaceae, Composi- 
tae, Ericaceae, Fagaceae, Rhamnaceae, and Rosaceae. Observations 
were made daily to determine if larvae were feeding and continuing to 
develop. This procedure was repeated for a group of fourth instars that 
had previously fed on greenleaf manzanita foliage. 

One day old egg masses collected from captured females were placed 
in the field. Prior to larval eclosion (ca. 10 days), egg masses were col- 
lected and transferred to Petri dishes. Adult parasitoids that emerged 
from eggs were collected and preserved. Larvae from a laboratory 
colony were periodically placed onto greenleaf manzanita branches in 
the field; those recovered several days later were reared individually to 
obtain parasitoids. 

StatistixO was used to perform all statistical analyses (Siegel 1992), 
following methods of Steel & Torrie (1980). Representative voucher 
specimens of all taxa have been deposited in the Maurice T. James En- 
tomological Collection at the Department of Entomology, Washington 
State University, Pullman, Washington. 


LIFE HISTORY 


Adult. Adult S. cervinaria flew at Hat Creek from early June to late July in 1991 and 
mid-May to late June in 1992 (Fig. 1). Adults have historically been collected in April, 
May, and June (Jones 1951), and early July (one specimen in the Essig Museum of Ento- 
mology, University of California Berkeley). Thus, based on these collection records and 
our data there appears to be one generation per year. 

During 1991, 160 males and 166 females were collected giving a capture sex ratio of 
nearly 1:1. In 1992, 102 males and 114 females were collected (1.0:1.1 ratio). Adults sur- 
vived up to 11 days in captivity. More than 94% of captured females produced eggs. All 
egg-producing females were fertile. Many S. cervinaria females attracted to the black light 


336 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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Fic. 1. Flight period of Synaxis cervinaria in 1991 (A) and 1992 (B). All adults at- 
tracted to a 15 watt black light were collected nightly from 2100—2300 h at the Forest In- 
sect Laboratory, USDA Forest Service, Hat Creek, California. Asterisks indicate adults not 
collected before 2 June 1991 nor between 26 May and | June 1992. 


| 


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showed little or no sign of wear (Fig. 2A), and had likely emerged within a few days of cap- 
ture. Virgin females placed in screen cages with adult males for 24 h produced viable eggs. 
These observations strongly suggest mating occurs soon after emergence from overwinter- 
ing pupae. Although mating in the field or laboratory was never observed, adults of at least 
some other geometrid species (e.g., Stamnodes animata |Pearsall]) are known to mate 
prior to flight (Furniss et al. 1988). An adult female collected in 1989 had seven sper- 
matophores within her bursa (K. Bolte, pers. comm. ). 

Female S. cervinaria use their papillae anales to locate a leaf edge. Eggs are deposited 
along the leaf edge, in a single row of several to about 30 (Fig. 2B). Under cramped artifi- 
cial conditions egg deposition also occurred along the edges of any available substrate 
(container lids, folded wax paper, cage frames, etc.), and they were often layered two or 
three deep. Eggs were always oriented with the micropylar end near the substrate edge. 


VOLUME 51, NUMBER 4 OT) 


FIG. 2. Synaxis cervinaria life stages. A, adult female. B, typical egg mass along leaf 
edge. C, fifth instar in a typical resting position on a greenleaf manzanita leaf. D, female 


pupa. 


Several egg masses were encountered in the field on greenleaf manzanita leaves and one 
egg mass was collected from a tobacco brush leaf at the Logan Lake site. 

More than 90% of the eggs produced by females in captivity were deposited after 2100 
h over a period of three or four nights. In 1991 a total of 159 captured females produced 
11,474 eggs for an average of 72 eggs per female (range: 0-230). In 1992, 94 captured fe- 
males produced 5207 eggs for an average of 55 eggs per female (range: 0-139). Some of 
these females may have deposited a portion of their eggs before flying to the black light. 
From a sample of 2110 eggs deposited by 30 females, 1846 produced larvae (87.5% hatch- 
ing rate). 


338 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Host suitability tests for Synaxis cervinaria larvae. No indicates that larvae 
do not feed and eventually starve to death; yes indicates that larvae feed and continue to 
grow and develop. 


Host family/species Feeding/frass 
Pinaceae 

Abies concolor (white fir) no 

Calocedrus decurrens (incense cedar) no 

Juniperus occidentalis (western juniper) no 

Pinus lambertiana (sugar pine) no 

P. ponderosa (ponderosa pine) no 

Pseudotsuga menziesii (Douglas-fir) no 
Compositae 

Artemesia tridentata (sagebrush) no 

Chrysothamnus viscidiflorus (yellow rabbitbrush) no 
Ericaceae 

Arctostaphylos patula (greenleaf manzanita) yes 
Fagaceae 

Chrysolepis sempervirens (bush chinquapin) yes 

Quercus kelloggii (California black oak) yes 
Rhamnaceae 

Ceanothus cordulatus (mountain whitethorn) yes 

C. integerrimus (deer brush) yes 

C. velutinus (tobacco brush) yes 
Rosaceae 

Cercocarpus betuloides (birch-leaf mountain-mahogany) yes 

C. ledifolius (curl-leaf mountain-mahogany) yes 

Prunus emarginata (bitter cherry) yes 

Purshia tridentata (antelope bitterbrush) yes 


Egg. Smooth, somewhat oblong (0.75 mm wide by 0.95 mm long) and remaining a 
pale yellowish-green for ca. 24 h following deposition. The eggs begin to tum a pinkish- 
rose color and then crimson after a second 24 h period (eggs that remain yellow are invi- 
able). About 12 h before hatch, the developing larva becomes visible through a translu- 
cent chorion, giving the eggs an overall reddish-gray appearance. Eggs hatch in 7-10 days 
depending on temperature. Parasitized eggs turn black prior to emergence of adult para- 
sitoids. 

First instar. Head capsule width (HCW) 0.45 + 0.02 mm, total body length (TBL) 
2.8—6.9 mm, stadium length (SL) 5 days (range: 4—7 days). Head capsule uniformly light 
brown; crimson dorsal and ventral stripes from pronotum to A9 dorsally, through A5 ven- 
trally, otherwise translucent until feeding commences, whereupon these areas take on a 
greenish hue; body lacks protuberances. 

Second instar. HCW 0.74 + 0.03 mm, TBL 6.1—13.3 mm, SL 6 days (range: 5—8 
days). Head capsule light brown with three sets of herring-bone patterns, more black than 
brown; crimson ventral region from cervix to A6 (first pair of prolegs); dorsum and lateral 
areas greenish-yellow due to food plant material visible through translucent integument; 
body lacks protuberances. 

Third instar. HCW 1.19 + 0.07 mm, TBL 11.9—21.1 mm, SL 6 days (range: 4—8 
days). Head capsule with more pronounced brown herring-bone pattern; fleshy dorsolat- 
eral lobes begin to appear on the metathoracic segment late in this stadium; body color 
varies to some extent; mottled black, brown, and crimson dorsally; ventrally and laterally 
an array of colors are exhibited including white, yellow, gray, brick red, crimson, brown, 
and black. 

Fourth instar. HCW 1.91 + 0.10 mm, TBL 18.2—25.5 mm, SL 8 days (range: 5-13 
days). Head capsule mottled/stippled brown, crimson, and white; fleshy tubercle at apex 


VOLUME 51, NUMBER 4 339 


TABLE 2. Greenleaf manzanita leaf consumption by Synaxis cervinaria larvae (n = 30). 
Total number leaves consumed determined by measuring the area of 528 mature leaves 
from 30 greenleaf manzanita branches to calculate a mean leaf area value (5.32 + 1.69 
cm?/leaf ). 


Percent of total 


lst—4th Instars 5th Instar foliage consumed 
Frass Leaf area Frass Leaf area lst—4th 5th Total number 
wt. (mg) consumed (cm2) wt. (mg) consumed (cm2) Instars Instar leaves consumed 


Mean 131 8.47 743 47.17 15.2 84.8 10.45 
SE 16 1.82 13 6.60 Dae 2.2, 1.82 


of clypeus; dorsolateral metathoracic fleshy lobes more pronounced; A4, A5, and A8 dor- 
sally with prominent paired chalazae; paired, dark colored ventrolateral and lighter col- 
ored ventromedial chalazae on Al; body coloration varies from crimson red to mottled 
gray with whitish hourglass patterns dorsally. These color variations are similar to those 
observed in both live and dead greenleaf manzanita twigs and branch stubs. 

Fifth instar. HCW 2.83 + 0.10 mm, TBL 24.3—35.2 mm, SL 14 days (range: 9-21 
days). Head capsule mottled/stippled light to dark brown, crimson, and white; fleshy tu- 
bercle at apex of clypeus; dorsolateral metathoracic fleshy lobes less pronounced; A4, A5, 
and A8 with prominent, paired chalazae dorsally, paired dark colored ventrolateral and 
lighter colored ventromedial chalazae on Al; body coloration and color patterns, as in in- 
star four, extremely variable. Stem/twig mimicry by cryptic morphology and behavior are 
pronounced in the fifth instar (Fig. 2C). 

Pupa. Obtect, mottled brown or tan, about 5 mm at the widest point and up to 19 
mm in length. Female genitalia span the 8th and 9th abdominal segments (Fig. 2D) 
whereas the male genitalia on the 9th segment only has a raised longitudinal border. Pu- 
pation occurs in leaf litter below host plants. 


BIOLOGICAL NOTES 


Larval behavior. Larvae developed normally on a variety of woody 
plants, but not on conifers or composites in the feeding trials (Table 1). 
Feeding tests on greenleaf manzanita showed that larvae (n = 30) con- 
sumed, on average, a total of 55.64 + 7.48 cm? of foliage or 10.45 + 1.82 
expanded leaves (Table 2). 

Neonate larvae emerge from eggs by using their mandibles to chew 
along a visible circular suture on the micropylar end of the egg, leaving 
a whitish-translucent shell with an apical exit hole. No parts of the 
chorion are consumed. There was no evidence of cannibalism for this or 
any subsequent instar, even under crowded conditions with no available 
food. Early instars feed by grazing the outer layers of cells from old and 
new foliage (greenleaf manzanita is a broadleaf evergreen that retains its 
leaves for more than one year), and although new foliage is preferred, 
larvae can develop normally on old foliage. Larval activity in late spring 
coincided with greenleaf manzanita leaf expansion. At the Logan Lake 
site, leaves first began to expand between 14—21 June in 1991 and 1-5 
June in 1992, about 10 days later than at Hat Creek (494 m lower in el- 
evation) where adults were surveyed. 


340 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


First instars released onto greenleaf manzanita branches in the field 
usually remained on or close to (within 15 cm) the leaf on which they 
were originally placed. Occasionally a larva would be found on a lower 
stem, possibly forced from an upper branch after disturbance by wind 
or predator. Unidentified species of ants (Formicidae) were ubiquitous 
and occasionally approached larvae which then dropped from their 
perch on a silken thread. Eight such encounters were observed in the 
field but none resulted in captured larvae. 

Prior to the first molt, the ventral area remains crimson but the dorsal 
stripe degenerates, and the dorsum and lateral areas become greenish- 
yellow. Laboratory-reared larvae were often observed in the evening 
suspended from silken threads (1.5—2.5 cm) or maintaining a character- 
istic stick-like appearance on the edge of a leaf (typical resting posture). 
The body is held in a straight and stiff position at a 30—45° angle to the 
substrate (usually a leaf or branch) to which the prolegs are attached, al- 
ways with a silk thread from the head to an attachment site on the sub- 
strate. Larval feeding was observed both day and night. 

Second instars feed by grazing on leaf surfaces and occasionally cre- 
ate a small hole in maturing leaves. Newly expanding leaves are some- 
times consumed by feeding along the leaf edge. Larvae released in the 
field tended to remain on or near the point of placement if left undis- 
turbed, similar to behavior observed for first instars. 

Third instars released in the field begin to move about on the fotees 
more so than the first and second instars, but they still remain relatively 
close to the point of release. A stick-like resting posture is maintained 
on or near foliage. Feeding usually occurs along leaf edges and whole 
leaf sections are removed. 

The significance of cryptic coloration in the first three instars is not 
well understood. They are exposed in the canopy and presumably would 
be vulnerable to bird predation. However, field exclosure experiments 
revealed that birds had no significant effect on the survival of instars 1—3 
(Valenti 1994). 

Stem mimicry and crypsis in the fourth instars is made possible by a 
combination of resting posture, morphology, and coloration. During the 
day, larvae in this stadium begin to move down and away from the 
crown. Typically, fourth instars released in the field would be found in a 
30—45° resting posture on a main stem below the crown. Larvae at this 
stage can consume entire leaves and feed nocturnally. 

The molt to the fifth instar begins with the late fourth instar terminat- 
ing its feeding and hanging by the prolegs with its head downward for 
several hours (the process described here is similar for the other in- 
stars). The cuticle splits at the vertex and the old head capsule is 
sloughed off by the thoracic legs. With undulating body movements, the 


VOLUME 51, NUMBER 4 34] 


old skin is forced up towards the prolegs. Once the exuvium is slipped 
over the first pair of prolegs (A6) the larva transfers its anterior end back 
up to the leaf or branch, secures a perch with the thoracic legs, and then 
slips its posterior end out of the remaining old cuticle. The prolegs are 
then firmly attached to the branch along with a silk thread near the 
head. Once initiated, the entire molting period lasts about five minutes. 
The larva maintains itself in a typical motionless, stick-like appearance 
while it hardens and darkens. The shriveled exuvium remains attached 
to the branch, often long after molting is complete. 

Fifth instars account for ca. 36% of the entire larval developmental 
period and inflict the most damage to greenleaf manzanita foliage (85% 
of the total amount of foliage consumed) (Table 2). They feed at night 
on entire leaves and remain in the lower crown or near the plant base 
during the day. 

Fourth and fifth instars avoid detection by visually searching preda- 
tors (e.g., birds) by resting during the day below the crown. Their re- 
semblance to stems and twigs and their resting posture make them very 
difficult to detect. Although we never observed significant defoliation in 
the field and there have been no outbreaks of S. cervinaria reported in 
the literature, caged larvae completely defoliated greenleaf manzanita 
plants, suggesting that this geometrid is strongly regulated at low densi- 
ties by natural enemies. Exclosure experiments in the field supported 
this supposition; in the absence of birds and ants larval survival in- 
creased nearly five-fold (Valenti 1994). 

Pupae. The fifth instar drops or crawls to the ground and burrows 
one to two cm beneath the leaf litter surface where it spins a loose co- 
coon attached to pieces of litter and detritus. After 4 days (range: 3-7 
days) the molting period is complete and a hardened and darkened pupa 
is formed. The pupa overwinters. The average weight of 30 pupae was 
233 + 22 mg. Adults emerged from pupae cold treated at 4°C for 
90-120 days. 

Natural enemies. Two species of wasps were reared from para- 
sitized S. cervinaria eggs: Trichogramma sp. (Hymenoptera: Trich- 
ogrammatidae) and Telenomus alsophilae Viereck (Hymenoptera: Sce- 
lionidae). Individual parasitized S. cervinaria eggs produced two to five 
individuals of Trichogramma sp. or a single individual of Telenomus al- 
sophilae. 

Several parasitoids were reared from S. cervinaria larvae (instars at- 
tacked are in parentheses): Campylochaeta sp. (Diptera: Tachinidae) 
(4,5), Aleiodes n. sp. (Hymenoptera: Braconidae) (3), Meteorus rubens 
(Nees) (Hymenoptera: Braconidae) (1-5), Dusona nigritibialis (Vier- 
eck) (Hymenoptera: Ichneumonidae) (3—5), Euplectrus sp. poss. plathy- 
penae Howard (Hymenoptera: Eulophidae) (1—5). 


342 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


In the field, a female Goniozus gracilicornis (Kieffer) (Hymenoptera: 
Bethylidae) was observed dragging a moribund third instar of S. cer- 
vinaria across a greenleaf manzanita leaf. It is unknown if it had para- 
lyzed the larva. On three occasions spiders where observed successfully 
attacking S. cervinaria larvae. These were Misumenops celer (Hentz) 
(Araneae: Thomisidae), Xysticus sp. (Araneae: Thomisidae), and 
Metaphidippus sp. (Araneae: Salticidae). 


ACKNOWLEDGMENTS 


Klaus B. Bolte (Crop Protection Division, Agriculture Canada, Ottawa, Ontario) kindly 
identified a series of Synaxis cervinaria during the early stages of this project. His taxo- 
nomic expertise is greatly appreciated, as is the assistance of the following taxonomists, all 
of whom contributed to this study by identifying natural enemies of S. cervinaria: L. Mas- 
ner (Scelionidae) (Crop Protection Division, Agriculture Canada, Ottawa, Ontario); R. W. 
Carlson (Ichneumonidae), P. M. Marsh (Braconidae), A. S. Menke (Bethylidae), M. E. 
Schauff (Eulophidae), D. L. Vincent (Trichogrammatidae), and N. E. Woodley (Ta- 
chinidae) (USDA-ARS, Systematic Entomology Laboratory, Beltsville, Maryland), and R. 
Crawford (Araneae) (The Burke Museum, University of Washington, Seattle, Washing- 
ton). Patrick Shea (USDA Forest Service) kindly provided facilities at the Hat Creek 
Forest Insect Laboratory. We are grateful to John J. Brown, Gary L. Piper, and William J. 
Turner for their constructive comments and suggestions on an earlier version of this man- 
uscript. We also thank Lawrence F. Gall, Charles V. Covell, Jr., and an anonymous re- 
viewer for their helpful suggestions. Funding for this study was made possible through a 
cooperative agreement between the Department of Entomology, Washington State Uni- 
versity, Pullman, Washington, and the USDA Forest Service, Pacific Southwest Forest and 
Range Research Station, Redding, California (No. PSW-89-0017CA). Additional funding 
was provided by the United States Environmental Protection Agency through a research 
grant awarded to Alan A. Berryman (Agreement No. R820465-01-1). 


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Received for publication 9 April 1996; revised and accepted 22 September 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1997, 344-357 


REVIEW OF THE NEW WORLD BAGISARINAE WITH 
DESCRIPTION OF TWO NEW SPECIES FROM THE 
SOUTHERN UNITED STATES (NOCTUIDAE) 


DOUGLAS C. FERGUSON 


Systematic Entomology Laboratory, Agricultural Research Service, USDA, c/o U.S. 
National Museum of Natural History, Washington, D.C. 20560, USA 


ABSTRACT. The Bagisarinae are discussed and characterized, including the Old 
World genus Xanthodes Guenée, 1852, which is considered to be congeneric, at least in 
part, with the New World Bagisara Walker, 1858. This possible synonymy is complicated 
by the type species of both genera being somewhat atypical. Described as new are Bagi- 
sara praecelsa, from Texas and northeastern Mexico, and Bagisara brouana, from 
Louisiana and Mississippi. An identification key is provided for the 19 described New 
World species, which occur mostly in North and Central America. Taxonomic changes in- 
clude two new synonymies, one new combination, one revised status, and removal of the 
unrelated Central American Xanthia patula Druce from the genus Bagisara. 


Additional key words: moths, taxonomy, Bagisara, Xanthodes, Malvaceae. 


The 17 described species of the genus Bagisara Walker comprise the 
New World component of the subfamily Bagisarinae (Crumb 1956:76, 
Poole 1989 vol. 1:154, Kitching & Rawlins, 1997). Nine of these are 
listed as occurring north of the Mexican border (Franclemont & Todd 
1983:134). Two new species described in this paper enlarge the genus to 
19 species and increase to 11 the number of species recorded from the 
United States. Two or three additional species from the neotropics are 
unidentified and probably undescribed. 

The Bagisarinae are a peculiar group with respect both to larval mor- 
phology and adult genitalia. Forbes (1954:170) treated them as one of 
his “isolated genera” within a broad concept of the Acronictinae. Fran- 
clemont and Todd (1983:134) regarded them as a tribe of the Aconti- 
inae, although Crumb (1956:4, 76) had earlier elevated the group to 
subfamily rank because of unusual larval features and proposed the 
name Bagisarinae. Poole (1989:154) followed Crumb and maintained 
subfamily rank for Bagisara, while keeping the closely related Old 
World genus Xanthodes Guenée in the subfamily Chloephorinae. Com- 
mon (1990:457) also included Xanthodes in the subfamily Chloephori- 
nae. Kitching and Rawlins (in press) include both Xanthodes and Bagis- 
ara in the Bagisarinae, a conclusion with which I agree. 

The larva lacks prolegs on abdominal segments three and four, a con- 
dition not unusual in Noctuidae, including acontiines, and the uniordi- 
nal crochets on each of the remaining prolegs are appendiculate, each 
bearing a large, subapical tooth. However, some Acontiinae, including 
Amyna Guenée, also have the subapical tooth (Gardner 1946:65—68). 
The SV group has two setae instead of one on abdominal segment 7, 


VOLUME 51, NUMBER 4 345 


agreeing in this respect only with those of Acronicta Ochsenheimer, 
Rivula Guenée, and the Agaristinae among known noctuid larvae 
(Crumb 1956:4, 76, pl. 4A, Godfrey 1987:550). The third segment of the 
labial palp in the larva is unusually long, at least three times as long as 
the basal segment. However, these morphological observations were 
based on few species. Crumb (1956) appears to have had larvae of only 
one species, Bagisara rectifascia (Grt.); and can we be sure that it was 
not the superficially similar, more common, but atypical B. repanda (F.)? 
I found no voucher specimens. 

The few recorded host plants are species of Malvaceae. The Old 
World species (of Xanthodes) also are reported to feed on Malvaceae, 
including cultivated cotton (Gossypium) and Hibiscus; and an Australian 
species, Xanthodes congenita (Hamp.), feeds on Brachychiton para- 
doxum Schott (Sterculiaceae) (Cacao family) (Common 1990:457). 

The male genitalia (Figs. 12-17) have a distinctive configuration, with 
the valves usually fused together on the mesoventral (saccular) margin 
so that they cannot be spread apart in dissection without damage or dis- 
tortion. The juxta is not recognizable. Each valve usually has 3—4 apices 
or preapical processes, these being the membranous apex, the rounded 
free distal end of an apparent costal sclerite, and one or two more slen- 
der, elongate processes that may represent the clasper or digitus. Not all 
processes are present in all species. Large, eversible coremata, opening 
laterally, reside in the bases of the valves of some species but are missing 
in D. brouana n. sp., rectifascia, and repanda. The eighth segment of the 
male abdomen has an elaborately modified, broom- or fan-shaped, usu- 
ally heavily setose (hairy) sternite in the eighth sternum, as well as a 
ringlike or U-shaped eighth tergite (Figs. 9-11). The females have a 
conspicuous, characteristic bulla seminalis that may be delicately orna- 
mented with encircling bands of fine, radiating, fanlike, sclerotic rods. 

The moths have smooth, cylindrical, upturned palpi that exceed the 
upper margin of the front to about the level of the basal antennal seg- 
ment. The antennae are simple, filiform, and minutely setose in both 
sexes. The legs have long, shaggy vestiture on the femora and tibiae in 
all species examined except B. repanda, in which the scaling is smooth. 
Bagisara repanda is also unusual in having a large, conspicuous patch of 
black-tipped scales on the front surface of the male foretibia. 

While American members of this complex have continued to be 
treated as a single genus in the Bagisarinae or Bagisarini, their similar 
Old World relatives have been referred in recent years to the 
Chloephorinae (e.g., Poole 1989, vol. 2:991, Common 1990:457). My 
observations during preparation of this paper convinced me that the 
Bagisarinae and at least part of the species in the Old World genus Xan- 
thodes belong not only to the same subfamily, but to the same genus. 


346 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-8. Bagisara species. 1, B. brouana, holotype. 2, B. brouana, 2 paratype, 4.2 mi. 
NE of Abita Springs, St. Tammany Parish, Louisiana, 7 June 1984, V. A. Brou. 3, B. 
brouana, ¢ paratype, Lizana, Harrison Co., Mississippi, 1 July 1991, R. Kergosien. 4, B. 
brouana, ° paratype, same data as for Fig. 3 but collected 19 July 1991. 5, B. praecelsa, 
holotype. 6, B. praecelsa, 2 paratype, Mt. View Acres, San Antonio, Texas, 9 September 
1971, A. & M. E. Blanchard. 7, B. gulnare 3, Ames, Iowa, 4 July 1964, W. S. Craig. 8, B. 
gulnare 2, Lacon, Illinois, 10 July 1967. 

Fics. 9-11. Bagisara species, 5 abdominal structures. 9, B. praecelsa. 8th abdominal 
segment: tergite (left), sternite (right). 10, B. gulnare, sternite of segment AS. 11, B. 
brouana. 8th abdominal segment: tergite (left), sternite (right). 


VOLUME 51, NUMBER 4 347 


Fics. 12-17. Bagisara species, 3 genitalia. 12, B. gulnare. 13, aedeagus of same spec- 
imen. 14, B. brouana. 15, aedeagus of same specimen. 16, B. praecelsa. 17, aedeagus of 
same specimen. 


348 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Similarity of larval characters was first noted by Crumb, based on com- 
parison of his observations with those of Gardner (1946, 1948) in India. 
However, the unique genitalia appear to have been neglected. Xan- 
thodes transversa Guenée, a colorful, well-known tropical Asian and 
Indo-Australian species that ranges north to Japan, is similar to Bagisara 
species in all essential details, including the complex genitalia of both 
sexes. The moths have the same kind of wing pattern, and the larvae of 
Xanthodes and Bagisara species have related food plants: mainly species 
of Malvaceae. 

I do not synonymize Bagisara to Xanthodes in this paper because 
more revisionary work is needed, especially on the Old World species. 
Indeed, the type species of both Bagisara and Xanthodes are somewhat 
atypical, leaving doubts as to what the limits of the genus or genera 
should be. The type species of Xanthodes Guenée, 1852, is Phalaena 
malvae Esper [1796], now known as Xanthodes albago (Fabricius 1794), 
of the Old World tropics to the Palearctic, including southern Europe. 
In X. albago the valves are not fused together along their outer saccular 
margins, although closely approximate, and the distinctive sclerites (ter- 
gite and sternite) as shown in Figs. 9, 10, are not developed. The 12 Old 
World species of Xanthodes, as listed by Poole (1989), appear more di- 
verse than their American counterparts. 

The type species of Bagisara Walker, 1858, is B. incidens Walker, 
which is a junior synonym of Bagisara repanda (Fabricius 1793), a wide- 
spread and generally abundant neotropical species that reaches the 
southeastern USA. It differs from other American species in its smooth 
vestiture, somewhat different genitalia (e.g., no long processes on valve), 
slightly angulate outer margin on the forewing, and in having a patch of 
specialized black scales on the foretibia. 

The following key should help in the identification of all species cur- 
rently assigned to Bagisara. One new combination and four changes in 
synonymy are made in the key, and Xanthia patula Druce (1898:486, pl. 
94, fig. 14), from southern Mexico, Guatemala, and Costa Rica, is re- 
moved from Bagisara, where it was placed by earlier authors. Its geni- 
talia are very different from those of the Bagisarinae and of a more con- 
ventional type. I could not place it to genus, but am sure that the species 
belongs somewhere in the large assemblage long known as the Am- 
phipyrinae but now probably within the expanded concept of the 
Hadeninae of recent authors (e.g., Kitching & Rawlins, in press). It has 
a rich golden orange-brown forewing with darker brown markings and 
two white spots in the reniform. Otherwise all species and their syn- 
onymy remain as listed by Poole (1989), and the reader is referred to 
that work for nomenclatural detail that is not repeated here. Poole’s cat- 
alogue also provides references to illustrations for about 12 of the 19 


VOLUME 51, NUMBER 4 349 


Fics. 18-21. Bagisara species, ° genitalia. 18, B. praecelsa, with modified scale 
patches of seventh sternum removed. 19, B. praecelsa, with modified scale patches in 
place. 20, B. gulnare. 21, B. brouana. 


350 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


species plus Xanthia patula Druce (although the figure cited for B. de- 
mura is misidentified, as mentioned later). To these may be added the 
colored illustrations of B. gulnare and B. rectifascia by Rings et al. 
(1992, pl. 13, figs. 20, 25). Most species are easily identified by differ- 
ences in size, color, wing pattern, and other features as used in the key. 


KEY TO NEARCTIC AND NEOTROPICAL SPECIES OF BAGISARA BASED 
ON SUPERFICIAL FEATURES 


1. Postmedial line of forewing sharply or abruptly angled on Rs or M, ............ 2 
1’. Postmedial line with a more rounded curve, not abruptly angled ............. i 
2. Forewing greenish, olivaceous, or brownish gray; transverse lines whitish or at 
least paler than ground color (if forewing brown, transverse lines bicolored 
whitish and orange); forewing with or without darkened patch having pro- 
nounced brassy sheen im!subbterminall area of forewing 20.) ee 5 
2’. Forewing reddish brown to gray brown, or pale with brown markings; transverse 
lines darker than ground color; forewing never with darkened patch with brassy 
sheen in-subterminal area’ ..-.) 0. eb pel ae eee 6 
3. Forewing greenish or olivaceous; transverse lines pale, not bicolored; forewing 
with transversely elongate patch in subterminal area that is darker than rest of 
wing and has brassy sheen; hindwing grayish (an unidentified species from Para- 
guay, probably undescribed, would key to couplet 4 except that it lacks the 
darkened subterminal area with! brassy:sleen)\ sc) = ee 4 
3°. Forewing usually reddish brown with transverse lines plain or bicolored; fore- 
wing without darkened patch with brassy sheen in subterminal area; hindwing 
reddish browns although paler tham forewing) = a...) ne ee eee 5 
4. Angle of subterminal line reaching outer margin, antemedial line angled holon 
costa; dark zone with brassy sheen between subterminal line and outer margin 
of forewing not strongly contrasting with rest of wing. Wing length 15-18 mm. 
MidwestemU:Suete 28 ees a. cok Gee eee gulnare (Stkr.) 
4’. Angle of subterminal line not reaching outer margin, antemedial line straight or 
only slightly bent upon approaching costa; dark zone with brassy scales between 
subterminal line and outer margin of forewing strongly contrasting with paler 
color of rest of wing (as in B. albicosta). Wing length 14-17 mm. Texas, Mexico 
SADE aa te he REO GRE ee Ac a, 2 Seo praecelsa, n. sp. 
5. Transverse lines of forewing bicolored, orange brown on proximal side, whitish 
or gray white on distal side. Wing length 16-19 mm. Central Mexico to 
Honduras 2 oa eg oe Sn. ste adil ae ee ene laverna (Druce) 
5’. Transverse lines of forewing plain, not bicolored. (Similar to B. laverna or more 
grayish, paler hindwing). Wing length 15 mm. Cuernavaca, Morelos, Mexico 
ioe lg Ae Seite Nie eed RP ee ro PCO A, aie ae a malacha (Druce) 
(Note: Two species are figured in the literature as B. malacha, and I assume that 
the original one (Druce 1889, pl. 28, fig. 14) is correct, although I have not seen 
specimens. The species shown by Draudt (1926, pl. 44, row a) is B. oula. Druce 
(ibid., p. 305) described the forewing as having “three narrow dark brown lines,” 
but his figure shows the usual pale lines of this group, perhaps with only a thin, 
dark edging. B. malacha appears close to B. laverna but smaller, and it is without 
the feature of the orange-brown margin on the transverse lines, which I describe 
as bicolored). 
6. Male forefemur with large tuft of long, reddish-brown scales, and male forewing 
with squarish black spot on costa between base and antemedial line (often re- 
duced, missing, or rubbed off); forewing otherwise pale with reddish-brown 
markings and a characteristic double longitudinal streak through angles in post- 
medial and subterminal lines. Wing length 12-13 mm. Texas and Arizona to 
AT@embina a0 0 ie ad lee GOR eed ori tc es re age mene Tr a are tristicta (Hamp.) 


VOLUME 51, NUMBER 4 351 


6’. 


10. 


10’. 


11. 


Wakes 


12. 


ie 


13. 


Ike 


14. 


Male forefemur without large scale tuft, and male forewing without black costal 

spot; forewing reddish brown to yellowish or gray brown, without a double or 

usually any streak through angle of postmedial and antemedial lines (except 
aarpecomasswnichihasonedark: streak)! 08.05 oo sd ee 7 
Forewing dark reddish brown or dusted with brown scales against a paler back- 
ground; angle of postmedial line less than 90°; forewing with dark longitudinal _ 
streak through angles of postmedial and subterminal lines; hindwing light reddish 
brown with thin, darker brown postmedial and subterminal lines, the former angled 

to mimic that of forewing (the only species with such a pattern on hindwing). 
iWimnedencthlo—l7 mm. (Guerrero, Mexico: 42. 3025... 5.0.5- graphicomas (Dyar) 


. Wings reddish brown to gray brown or yellowish; angle of postmedial line acute 


or obtuse; forewing without dark longitudinal streak through angles of postmedial 

and subterminal lines; hindwing without markings. Wing length variable. 

ON IGIESIOIREAG! « 6 tu yn See cata EE Ce LAU ons he aS ck) oa and ae ee ee REO rao ae 8 
Both wings bright reddish brown, hindwings paler but almost uniformly colored; 
forewing with well-marked, clearly defined, regular, dark-brown, transverse lines; 
angle in postmedial line of forewing usually 90° or less (acute) ................ 9 
Forewing light violaceous gray brown to yellowish brown; hindwing whitish, pro- 
gressively shading to pale brownish distally; forewing with thin, delicate, regular, 
brown transverse lines; postmedial line acutely or obtusely angled ............ 10 
VyinelenethelS=19imm! Orizaba, Veracruz ..........5250 05. ochracea (Schaus) 


. Wing length 15 mm. SW U‘S. to Mizantla and Guadalajara, Mexico .. demura Dyar 


(Note: B. ochracea and B. demura appear exactly alike except for size, but only 
females were available during preparation of key. Bagisara ochracea (Schaus 
1906) is a new combination. It was described as Trileuca ochracea Schaus and 
referred to the genus Schinia by Poole (1989). Bagisara xan Dyar, 1913, is a 
junior synonym of Trileuca ochracea Schaus, and this is a new synonymy. These 
changes are based on reexamination of the types in the U.S. National Museum 
of Natural History. Draudt’s figure (1926, pl. 44, row b) of B. demura is not that 
species but is B. anotla; and his figure of anotla (1926, pl. 44, row b) is B. ochracea. 
An unidentified species similar to B. ochracea, but with more oblique lines, will 
key out here. It has been taken in southern Minas Gerais, Brasil (USNM)). 
Forewing light gray brown with faint violaceous tint; angle of postmedial line 
often less than 90°; antemedial line usually angled before costa. SW U.S., 


lee Ome REA OTIS) eh Wa ae es SRE Warn dae Bee oula Dyar 
Forewing light yellowish brown; angle of postmedial line about 90°; antemedial 

line usually bent but not angled before costa. SW U.S., Mexico ....... buxea (Grt.) 
Transverse lines of forewing whitish, or at least paler than ground color ........ 12 
Transverse lines of forewing brown to blackish, darker than ground color, 

Ramm inilesenomemtuinclyidistinctt), Sl away Bade HES Lyte Bt es Soh A 16 


Male foretibia and forefemur with large, dense, black-tipped scale tufts; outer 

margin of forewing slightly angulate; small, wing length 10-12 mm. SE U.S., 
RVestaindicsjandeMexico to: Paraguay) 4/52 as PO ri repanda (F.) 
Male foretibia and forefemur without large scale tufts; outer margin of forewing 

Pan cnlate vying length) variables: sh. ok PS ane ee See Se 13 
Antemedial line of forewing farther from base at costa than at inner margin. SW 

WR SRLOMVEMEAUIC late ireiee (ini Niel hatter Pence leben ie Soc! Sees albicosta Schaus 
Note: B. albicosta is the only species in which antemedial line of forewing is 

slanted in reverse direction from what is usual. Coloring of forewing closely 

similar to that of B. praecelsa, including even the dark subterminal area with 

brassy sheen, but course of lines is different). 

All transverse lines of forewing approximately erect from inner margin and sub- 


Poetics lemeiieyl Silos] MA ante Cpe Nese are eNO tes Ramee ay F APT WS seek ee Ohh rena ate caiits 14 
Wings dark brown; lines pale but faint. Small, wing length 11 mm. Guyana, 
(OG INMGey MAT Wis Reena ent o2 Meete Peete AS oe eA OERRS REO TAGs AP aRRES 18 Cem eA a obscura Hamp. 


(Note: B. obscura has the appearance of a very dark-suffused B. repanda, but 
the outer margin of the forewing is evenly rounded, not angulate. The wing 
shape and pattern, although indistinct, appear much as in B. rectifascia). 


352 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


14’. Wings light brown with distinct, pale (or partly pale, partly darker) transverse 
lines Vines emethy 2214: mamaria a eee aera ne ina, Se 15 
15. Forewing with inner margin straight or slightly convex; outer margins evenly 

rounded, tornus rounded; hindwing normal, not appearing produced or unusually 

triangular. U.S. to Nicaragua and Costa Rica (may not all be same species) 

jp cluncse ber einidieah BO be Pupite Sasi a uecnte  f Ad a han | ail ee ee rectifascia (Grt.) 

15’. Forewing with slightly concave inner margin and relatively sharply angled tornus; 

outer margins of both wings not entirely rounded but with places where they are 

straight or very slightly concave; hindwing appearing more triangular, with anal 

angle about 90° and not or hardly rounded. SW U.S. to Costa Rica . pacifica Schaus 
16. General wing coloring very dark, blackish, lines present but obscure. Wing 

length 13—15 mm iouisianas Mississippil a5. 4 4a. aaa eee brouana, n. sp. 
16’. Much paler, pale brown to reddish brown. Wing length 13-17 mm. Neotropical Wa 
17. Very pale brown with thin, regular, erect, subparallel darker brown lines on 


>) 


forewing. Wing length 12-15 mm. Panama, Brasil, Venezuela ... paulensis (Schaus) 
17’. Light reddish-brown moths with smudgy markings and lines often indistinct. 
Guatemala to Panama s.2 28h. 2 oe en ee ee eee 18 


18. Deep reddish-brown suffusion across much of forewing except costal area, and 
patch of blackish to dark-gray scales in median space near inner margin; hindwing 
whitish toward base, shading to yellowish brown distally. Small, wing length 13-15 
mn (@watennalasele amcarncues: 4-5. 5.) eee cen ee avangareza Schaus 
18’. Forewing without dark shades except some deep reddish brown near outer 
margin; hindwing bright orange brown, not or hardly paler toward base. Larger, 
wing length 16-17 mm. Mexico, Guatemala, Panama ............... anotla Dyar 
(Note: Bagisara lulua Schaus, 1921 is a junior synonym of B. anotla Dyar, 1914, 
new synonymy, and the latter is a separate species, revised status, not a 
synonym of B. demura Dyar, 1913, as indicated by Poole (1989:154). This is 
based on a reexamination of the Schaus and Dyar types in the U. S. National 
Museum, but few specimens are available and most are females. The figure of 
B. lulua by Draudt (1926, pl. 44, row b) is a good likeness of B. anotla). 


Bagisara praecelsa Ferguson, new species 
(Figs. 5, 6, 9, 16, 17, 18, 19) 


Diagnosis. This species resembles Bagisara gulnare (Strecker) and was misidentified 
as that species in collections. However, the two apparently are not sympatric, as the true 
B. gulnare is known from Ohio, Michigan, Illinois and Iowa, whereas B. praecelsa occurs 
in Texas and northern Mexico. They are easily distinguished by differences in the forewing 
pattern. The antemedial line is straight or only slightly incurved near the costa in prae- 
celsa, sharply angled near the costa in gulnare. Also, in praecelsa, the angles in the post- 
medial and subterminal lines are offset relative to each other; the outermost point of the 
postmedial is no farther out than the point at which the subterminal meets the costa, and 
the outermost point of the subterminal usually stops short of the outer margin. In gulnare, 
the angulate part of the postmedial fits within that of the subterminal with almost perfect 
symmetry, the point of the postmedial surpasses the point where the subterminal meets 
the costa, and the outermost point of the subterminal reaches the outer margin. In prae- 
celsa, the contrasting zone of metallic scales between the submarginal band and the outer 
margin is larger as a result of the submarginal band being more deeply incurved, and it is 
deep metallic red brown rather than gold colored. Although males of both species differ 
from all other U.S. Bagisara in having a distinct patch of erect scales on the hindwing be- 
tween the second and third anal veins about a third of the way out from the base, these 
scales are yellow in praecelsa and light grayish brown in gulnare. The complicated geni- 
talia of praecelsa are similar to those of gulnare in both sexes, but differ in the shape and 
proportions of many parts. 

Further description. Antenna simple in both sexes; male palp brushlike, tufted with 
stiff, hairlike scales on ventromesial surface; alula-like structure (posterior to base of hind- 
wing on each side) bearing an expandable tuft of long, yellow-brown scales matching in 


VOLUME 51, NUMBER 4 S15 


color the erect scales on hindwing and transverse dorsal intersegmental bands on abdomi- 
nal segments 5—8; tegula short compared to that of at least some other species of Bagi- 
sara, not reaching base of abdomen; all foregoing features common to gulnare and prae- 
celsa, except that sex scales in praecelsa are bright-yellow instead of gray-brown. Forewing 
light, lustrous, olivaceous gray, finely dusted with white scales, traversed by three thin, 
clearly defined transverse lines, and with a submarginal, lunate, metallic, dark coppery- 
brown patch about as wide as minimal distance between antemedial and postmedial lines 
(narrower in gulnare). Other forewing markings as described in diagnosis. Hindwing gray 
brown, often slightly darker toward outer margin, and with a terminal series of vague 
whitish dots or wedges between vein endings (rather than the continuous, slightly sinuous, 
whitish terminal band apparent in fresh specimens of gulnare). Length of forewing: holo- 
type, 16 mm; other dd, 14-17 mm (n = 25); 9°, 16-17 mm (n = 7). 

Male genitalia (Figs. 9, 16, 17). Differing from those of Bagisara gulnare (Figs. 10, 
12, 13) most obviously as follows: overall shape of valve and its everted corema more slen- 
der, and corema with hair tufts only half the size of those in gulnare (cut off in Fig. 12); 
valve with rounded end of costal lobe produced beyond end of most mesial of the slender, 
bladelike, preapical, valvular processes; vesica with two sclerites of nearly equal size (un- 
equal in gulnare), and a diverticulum smaller than that of gulnare. Sclerites of eighth ter- 
gum and sternum also differ, as illustrated. 

Female genitalia (Figs. 18, 19). Similar to those of Bagisara gulnare (Fig. 20) in 
most respects, but easily distinguished by the smaller, less persistent, paired scale tufts on 
sternum 7. These tufts are large, dark colored, and difficult to remove in gulnare (Fig. 20); 
smaller, paler, and more nearly deciduous in praecelsa. Also, in praecelsa, posterior mar- 
gin of sternum 7 relatively shallowly emarginate at ostium; outer two of four needlelike 
sclerites arising from ostium being one-third to four-fifths as long as middle pair; corpus 
bursae at juncture with ductus bursae bearing a relatively prominent sclerite marked with 
a fanlike, radiating pattern of ridges. 

Types. Holotype 4, Fort Davis, Jeff Davis Co., Texas, 11 July 1969, A. and M. E. 
Blanchard. Paratypes: 6 dd, same locality and collectors, 30 July 1964, 25 June 1965, 11 
June 1969, 28 August 1970; 1 °, same locality, 19 August 1984, E. Knudson; 2 4d, Mount 
Locke, 6700’, Davis Mountains, Texas, 10 June 1969, A. and M. E. Blanchard; 4 dd, 1 8, 
Alpine, Brewster Co., Texas, 10 June 1969, 2 August 1964, 6 Sept. 1964, 10 September 
1963, same collectors; 1 d, Bear Canyon, Guadalupe Mountains, Texas, 4 September 1969, 
same collectors; 2 dd, McKittrick Canyon, Guadalupe Mountains, Texas, 29 August 1967, 
same collectors; 1 °, Sierra Diablo Wildlife Management Area, 6000’, Culberson Co., 
Texas, 5 June 1969, same collectors; 2 dd, Junction, Kimble Co., Texas, 24 August 1973, 
same collectors; 3 dd, 1 2°, Mt. View Acres, San Antonio, Texas, 30 August 1973, 9 Sept. 
1971, same collectors; 1 °, Kerrville, Texas, June 1919; 1 5, San Benito, Texas, 16—23 
March [incorrect date?]; 1 5, Bentsen-Rio Grande Valley State Park, Hidalgo Co., Texas, 
27 May 1982, E. C. Knudson; 6 dd, 2 22, Limpia Canyon, Davis Mountains, Jeff Davis 
County, Texas, 4920’, 30° 17.4’ N, 103° 36.6’ W, 9 August 1991, E. H. Metzler; 1 2, 3 mi. 
E Galeana, 5000’, Nuevo Leon, Mexico, 7-9 August 1963, [W. D.] Duckworth and [D. R.] 
Davis. Holotype and most paratypes in collection of U. S. National Museum of Natural 
History; some paratypes returned to E. C. Knudson, E. H. Metzler, and deposited in other 
museum collections; namely, the Canadian National Collection, Ottawa; American Mu- 
seum of Natural History, New York; Carnegie Museum, Pittsburgh; Cornell University, 
Ithaca, New York, and The Natural History Museum, London. 

Distribution. This species has a wide distribution within Texas, as the above listed 
localities indicate. Otherwise, I have seen it only from Mexico, where the one paratype 
was collected about 95 km south of Monterrey and 240 km from the U.S. border. 

Early stages. Undescribed. The larva of B. gulnare feeds on foliage of glade mallow, 
Napaea dioica L. (Malvaceae). One specimen was reared from a larva found on this plant 
in remnant wet prairie in Pickaway County, Ohio by E. Metzler, and the moth is regarded 
as an endangered species in Ohio (Rings et al. 1992:71). The larva is green with a vague, 
yellowish lateral stripe. 

Remarks. Bagisara laverna (Druce), of Mexico and Central America, is the species 
most similar to B. gulnare and B. praecelsa in size, color, and pattern. Bagisara albicosta 


304 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Schaus, although nearly identical to B. praecelsa in size and especially coloring, has differ- 
ent transverse lines (see key). If the apparent allopatry of praecelsa and gulnare proves to 
be real, then these two species may be distinguished by locality label. I examined speci- 
mens of B. gulnare from four states, as follows: OHIO: 30 mi. S of Columbus (emerged 12 
Aug. 1989, 1 specimen in Metzler coll.). MICHIGAN: Berrien County (26 June-14 July, 3 
specimens, J. H. Newman, in Metzler coll.). ILLINOIS: Champaign; Elgin; Lacon; Peo- 
ria; Putnam Co.; Quincy; Urbana (9 June—1 Sept., 73 specimens in Illinois Natural History 
Survey collection and U. S. National Museum of Natural History). IOWA: Ames; Milford; 
Soldier (4 June—2 Sept., 8 specimens in U.S. National Museum of Natural History). Ba- 
gisara gulnare has also been recorded from Pennsylvania (Tietz [1952]:83; Forbes 
1954:264), but this needs verification. Neither gulnare nor praecelsa appears to have been 
identified or recorded in the literature from any of the intervening states such as Missouri, 
Mississippi, Arkansas, Kansas, and Oklahoma, although doubtless there are records that I 
have not seen. 


Bagisara brouana Ferguson, new species 
(Figs. 1-4, 11, 14, 15, 21) 


Diagnosis. A large, distinctive, dark-brown species known only from St. Tammany 
and Tangipahoa Parishes, Louisiana, where many have been collected over a period of 
years by the collector for whom it is named, and from Hancock and Harrison counties, 
Mississippi, where it was collected by R. Kergosien. The forewing pattern of three narrow, 
subparallel, transverse bands is generally similar to those of B. rectifascia and B. repanda, 
but the bands are largely blackish on a dark-brown background, not pale on a light-brown 
background as in those species. The bands are often inconspicuous because of the overall 
dark suffusion. Bagisara brouana is unique in being sexually dichromatic with respect to 
wing color, the males being distinctly lighter on both fore- and hindwings. In our fauna it 
is most closely related to B. rectifascia, and, like that species, has unmodified vestiture on 
the male foreleg and an evenly convex outer margin on the forewing. Bagisara repanda 
differs in having a conspicuous patch of long black scales on the male front femur and a 
somewhat angulate outer margin on the forewing. | 

Further description. Male foreleg unmodified, without unusually long or special- 
ized scales, and without a femoral tuft of black scales. Antennae and other external struc- 
tures as in related species. Forewing of both sexes with outer margin evenly convex, not 
angulate near middle as in B. repanda, dark reddish brown (blackish) with a violet irides- 
cence in fresh specimens; discal spot transversely oblong, blackish, diffuse; the three trans- 
verse bands narrow, blackish (sometimes partly shadowed by a pale shade), almost erect 
relative to inner margin, and in nearly the same positions as the pale bands of B. rectifas- 
cia; antemedial band straight or slightly concave, about one-third of the way out from 
base; postmedial band just beyond discal spot, slightly concave except toward costa, where 
it curves basad; subterminal band meeting inner margin at or near tornal angle in B. 
brouana and B. rectifascia (midway between postmedial band and tornal angle in B. 
repanda); an incomplete terminal row of small black dots usually present; males often with 
space between postmedial and subterminal bands occupied by a slightly paler, violaceous- 
brown shade that may form an almost complete pale band (as in holotype) or be confined 
to a patch near inner margin; fringe concolorous with wing or darker. Hindwing sexually 
dimorphic in shape and coloring; that of male with outer margin often nearly straight be- 
tween M, and first anal fold; light yellowish brown, glossy, with dusky shading near inner 
margin and, in most fresh males, with gray, elongate, wedge-shaped rays toward outer 
margin and between veins; female hindwing with outer margin evenly and roundly con- 
vex, and the color uniformly dusky brown, nearly as dark as forewing; discal spot wanting 
in male, nearly so in female; fringes concolorous. Scales of thorax concolorous with 
forewing, of abdomen concolorous with hindwing. Underside pale, dusted with darker 
gray-brown scales, with discal spots developed on both wings, and with rounded, evenly con- 
vex, parallel, dark-brown postmedial bands variably developed on both wings, poorly so in 
males, often half-developed on forewing and fully developed on hindwing in females. Length 
of forewing: holotype, 13.5 mm; other 53, 13-15 mm (n = 18); 9°, 12-15 mm (n = Sy) 


VOLUME 51, NUMBER 4 355 


Male genitalia (Figs. 14, 15). Similar to those of other species of Bagisara examined 
inasmuch as the ventral margins of the valves are fused together so that the valves cannot 
be spread apart without damage and distortion. Genitalia most closely resemble those of 
B. rectifascia, but are greatly elongated to 1.5 times length from saccus to uncus, with all 
components proportionately elongated, including aedeagus. The complex eighth sternite 
(Fig. 11) similar in form but also elongated, its posterolateral apices not flared outwardly 
as much or as abruptly as those of B. rectifascia. Male genitalia differ in shape of virtually 
all parts from those of B. repanda and B. buxea. Eighth sternite and tergite are both com- 
plex in this genus, but in B. brouana and B. rectifascia are especially conspicuous; eighth 
sternum of these species bears a partly sclerotized structure (sternite) forming an elon- 
gated, funnel-shaped structure with a large, medial, posteriorly directed tuft, probably the 
corema associated with a scent gland; eighth sternite of B. repanda with a similarly situ- 
ated but smaller tuft, comprised mainly of long, spatulate scales. Bagisara brouana, recti- 
fascia, and repanda all lack the coremata arising from the bases of the valves in B. buxea, 
which clearly is not a closely related species. 

Female genitalia (Fig. 21). As illustrated. 

Types. Holotype 4, 4.2 mi. NE of Abita Springs, Sec. 24, T6 SR12E, St. Tammany 
Parish, Louisiana, 8 August 1983, V. A. Brou. Paratypes: 17 dd, same locality and collector, 
4, 11, May, 6, 11, 18, 19, 23 June, 2, 7, 27 July, 8, 9, 11, 13 August, 2 September 1983; 17 
33, same locality and collector, 5, 7, 14, 22, 29, 30 May, 7, 9, 10, 14, 21, 18 June, 21, 22 
July, 8, 30 August, 1 September 1984; 10 22, same locality and collector, 22 May, 6, 9, June, 
1, 16, 26 July, 1, 8, 14, 20 August 1984; 15 29, same locality and collector, 26, 29 April, 13, 
22, 28 May, 3, 6, 7, 20, 22 June, 18, 19 July, 1 August 1984; 2 °°, Fluker, Tangipahoa 
Parish, Louisiana, 12 May 1978, V. A. Brou; 30 33, 10 9°, Lizana, Harrison County, Missis- 
sippi, 16 June—17 August 1991, R. Kergosien; 6 dd, 7 22, Long Beach, Harrison County, 
Mississippi, 30 June—20 August 1991, R. Kergosien; 1 5, Blk. Crk. near George-Jac|kson] 
Co. line, Jackson County, Mississippi, 1 August 1991, R. Kergosien; 1 2, Pass Christian, 
Harrison County, Mississippi, 19 July 1979, R. Kergosien; 1 °, Bay St. Louis, Hancock 
County, Mississippi, 16 July 1979, R. Kergosien. The Louisiana specimens were all col- 
lected in light traps using mercury vapor lamps and ultraviolet fluorescent tubes. Holotype 
and some paratypes in collection of U.S. National Museum of Natural History; remaining 
paratypes mostly in collection of V. A. Brou, Bryant Mather, and R. Kergosien, but some 
will be distributed to other collections as mentioned under B. praecelsa. Fifteen additional 
specimens from the type locality were examined but not labelled as paratypes because of 
their poor condition. 

Distribution. Known only from the Louisiana and Mississippi localities listed above. 

Early stages. Unknown. Bagisara rectifascia has a slender green semilooper larva, 
with the first two pairs of prolegs missing, reported on Hibiscus lasiocarpus Cav. and Mal- 
vaviscus drummondii T. & G. (both Malvaceae) in Texas (Crumb 1956:77). Larva of B. 
buxea reported on a species of Sphaeralcea, also in the Malvaceae (Comstock & Dammers 
1935:138). 

Remarks. A curious feature of this species is its restricted distribution. Intensive col- 
lecting in the southern States in recent years has failed to reveal its presence anywhere 
outside of the three coastal counties of Mississippi and two nearby parishes of Louisiana, 
as far as I am aware. However, nearly 200 specimens collected by Brou in two seasons at 
the type locality indicate a large and thriving local population. It may be a specialized 
feeder on one genus or even one species of plant that also has a limited distribution. The 
closely related B. rectifascia has a wide distribution in the eastern U.S. but is not common 
in collections. Bagisara repanda is sometimes abundant in the Southeast, particularly in 
Florida, and throughout the Caribbean Region and much of the American tropics. The 
only other species of Bagisara regularly present in the U.S. east of Texas and the Great 
Plains is gulnare (Strecker) (Figs. 7, 8), a similarly large but conspicuously different spe- 
cies with an olive-green forewing having a brassy sheen and oblique, silvery-white bands. 
It was further discussed above under B. praecelsa. Bagisara buxea has been reported as 
far north as Wisconsin (type of delicia (Dyar), a junior synonym of buxea), perhaps as a va- 
grant from the South or with a false locality label. Bagisara gulnare and B. rectifascia were 
illustrated in color by Rings et al. (1992, pl. 13, figs. 20, 25), and Bagisara repanda (as 


356 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Atethmia subusta Hbn.) and B. rectifascia were illustrated (although not very well) by 
Holland (1903, pl. 27, figs. 4, 5). 

A summary of Louisiana Bagisara records to the end of 1995 sent by V. A. Brou shows 
an almost continuous flight period for B. brouana from about 6 April to 19 September, 
with major peaks in early and late June, lesser peaks in late May, late July, and early to mid 
August, and an abrupt decline in September. This would seem to reveal three or more ex- 
tended, overlapping generations. Comparable data for Bagisara rectifascia, but with far 
fewer records, show a similar pattern. In contrast, the many records for B. repanda are 
mostly clustered late in the season, from late August into November, with the greatest 
number in September. This is the classic pattern of a seasonal immigrant from the tropics 
that may not overwinter in the United States. 


ACKNOWLEDGMENTS 


Itisa pleasure to name Bagisara brouana after its original discoverer, Vernon A. Brou 
of Abita Springs, Louisiana, who brought the species to my attention and who provided all 
the known specimens except those subsequently sent from Mississippi by Bryant Mather. 
I thank E. C. Knudson (Bellaire, Texas), B. Mather (Jackson, Mississippi), and G. L. God- 
frey (formerly of the Illinois Natural History Survey, Urbana, Illinois) for the loan of ma- 
terial; E. H. Metzler (Columbus, Ohio) for host and distribution data; and R. W. Poole 
(Rockville, Maryland) for comments and advice on noctuids. Bagisara praecelsa is de- 
scribed mostly from material collected by the late André Blanchard and donated to the 
U.S. National Collection. I thank R. W. Hodges and J. Pakaluk of the Systematic Ento- 
mology Laboratory; J. G. Franclemont, Ithaca, New York; E. H. Metzler, Columbus, Ohio; 
J. S. Miller, American Museum of Natural History, New York; and D. F. Schweitzer of Port 
Norris, New Jersey for reviewing the paper. I prepared all illustrations. 


LITERATURE CITED 


COMMON, I. F. B. 1990. Moths of Australia. E. J. Brill, Leiden. 535 Pp: 129 ae 32 col. 

ls. 

Ooueeee J. A. & C. DAMMERS. 1935. Notes on the early stages of three butterflies and 
six moths from California. Bull. So. Calif. Acad. Sci. 34:120—142. 

CRUMB, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull. 1135. 
Washington, D.C. 356 pp., illus. 

DraubT, M. 1926. In Seitz, A. 1919-1944. Die Gross-Schmetterlinge der Erde. 
Abteilung II. Amerikanischen Faunengebietes. Band 7. Eulenartige Nachtfalter. Al- 
fred Kernen, Stuttgart. 508 pp., 96 pls. 

Druce, H. 1889. In Godman, F. D. & O. Salvin, 1881-1891. Biologia Centrali-Ameri- 
cana; or Contributions to the Knowledge of the Fauna of Mexico and Central Amer- 
ica. Zoology. Lepidoptera. Heterocera by H. Druce. Vol. 1. Taylor and Francis, Lon- 
don. 490 pp., pls. 1-64. 

. 1898. In Godman, F. D. & O. Salvin, 1891-1900. Ibid., vol. 2. 692 pp., pls. 
65— 101. 

FORBES, W. T. M. 1954. The Lepidoptera of New York and neighboring states, pt. 3, 
Noctuidae. Cornell Univ. Agric. Expt. Sta. Mem. 329. 433 pp. 

FRANCLEMONT, J. G. & E. L. Topp. 1983. Noctuidae, pp. 120-159. In R.W. Hodges et 
al. (eds.), Check list of the Lepidoptera of America north of Mexico. E. W. Classey 
Ltd. and The Wedge Entomol. Research Foundation, London. xxiv + 284 pp. 

GARDNER, J. C. M. 1941. Immature stages of Indian pe een 2. Noctuidae, Hypsi- 
dae. Indian Forest Records, Entomology 6:253—298, 2 pls. 

. 1946. On larvae of the Noctuidae (Lepidoptera)—1. Trans. Roy. Entomol. Soc. 

London 96:61—72. 

. 1948. On larvae of the Noctuidae (Lepidoptera)—IV. Trans. Roy. Entomol. Soc. 
London 99:291—318. 

Goprrey, G. L. 1987. Noctuidae, pp. 549-578. In F.W. Stehr et al. (eds.), Immature in- 
sects. Kendall/Hunt Publishing Co., Dubuque, Iowa. xiv + 754 pp., illus. 


VOLUME 51, NUMBER 4 35 


HOLLAND, W. J. 1903. The moth book. Doubleday, Page & Ce., New York. 479 pp., 48 
col. pls. 

athe. I. A. & J. E. RAWLINS. In press. Chapter 19, The Noctuidae. In N. Kristensen 
(ed.), Lepidoptera. Handbook of Zoology, v. 4. 

POOLE, R. W. 1989. Noctuidae. Lepidopterorum Catalogus (new series), fasc. 118. E. J. 
Brill/Flora & Fauna Publications, Leiden and New York. xii + 1314 pp. in 3 vols. 

RINGS, R. W., E. H. METZLER, F. J. ARNOLD & D. H. Harris. 1992. The owlet moths of 
Ohio, order Lepidoptera, Family Noctuidae. Ohio Biol. Surv. Bull., New Series, 9(2). 
vi + 219 pp., 16 pls. 

TIETZ, H. M. [1952]. The Lepidoptera of Pennsylvania. Pennsylvania State College, 
Agric. Expt. Sta., State College, Pennsylvania. xii + 194 pp. 


Received for publication 17 January 1996; revised and accepted 12 February 1997. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
51(4), 1997, 358-359 


EUREMA DINA LEUCE (PIERIDAE) FEEDS ON A 
LACTIFEROUS HOSTPLANT IN EASTERN BRAZIL 


Additional keywords: feeding behavior, Leguminosae, Mimosa, oviposition preference. 


The pantropical genus Eurema (Pieridae) includes 37 neotropical species (DeVries 
1987) whose larvae feed primarily on plants in the Fabaceae, Caesalpinaceae, and Mi- 
mosaceae (Clark & Dickson 1965, DeVries 1985, 1987, Jones & Rienks 1987, Jones et al. 
1987). In contrast to most species in the genus, the larvae of Eurema dina westwoodi Bois- 
duval utilize two species of Picramnia (Simaroubaceae) in Costa Rica (DeVries 1985, 
1987), and those of E. hecabe phoebus Butler are reported to feed on Euphorbiaceae in 
Australia (Jones et al. 1987). Brown (1992) also noted that larvae of E. elathea Cramer oc- 
casionally feed on Apocynaceae, but stressed that this species prefers legumes. 

We studied E. dina leuce Boisduval in an open area and along a forest edge at the Lin- 
hares Forest Reserve (19°8’S, 40°3’W), Espirito Santo, Brazil in July 1995. We observed 
two females ovipositing on leaves of Mimosa laticifera Rizzini and Mattos (Mimosaceae), a 
species with lactiferous leaves, a rare trait within the Leguminosae. The 2—4 pairs of large 
(2-3 cm diam.) leaflets of this plant present secondary venation confluent with the mar- 
ginal nerve and exude droplets of latex when leaf ducts are damaged. Mimosa laticifera is 
distributed throughout eastern Brazil southward from 7°S (Barneby 1985) and occurs as a 
shrub (0.1—1.0 m tall) at the study site. As far as we know, this is the first documented use 
of a latescent plant for a New World Eurema. 

To test whether E. dina leuce showed habitat preference for oviposition, we sampled 
eggs and larvae on 150 plants of M. laticifera in an open area and on 60 additional plants 
along a neighboring forest edge. Oviposition site preference was evaluated using the pro- 
portion of plants with early stages of E. dina leuce in each area. To estimate host-plant 
density, we established 17 plots of 64 m? in each area and counted the number of M. lati- 
cifera in 5 quadrats of 0.25 m2 placed sistematically in each plot (one at the center and 
four at the corners). 

We found a total of 39 eggs on the leaves of 23 of the 210 M. laticifera plants examined. 
We also encountered two first instar and two fourth instar larvae on these and two addi- 
tional plants which lacked eggs. Plants with eggs averaged 17.9 cm tall (SD = 7.1, n = 23) 
and did not include plants in the larger size categories. Field collected eggs were white 
and spindle-shaped and eclosed 3.4 days (SD = 0.9, n = 14) after oviposition (data for eggs — 
with known time of oviposition). Sixty percent of the eggs (n = 39) were on new leaves and 
40% on mature leaves. Most were found singly on plants (72%) usually on the underside 
of leaves (87%). Eggs censused in the field showed high mortality rates: 51% of the 39 
eggs disappeared before hatching and an additional 13% turned black without hatching, 
apparently due to parasitism. 

Eurema dina leuce strongly preferred plants growing along the forest edge. As stressed 
by Stanton (1982), selectivity is manifested when certain resource types are utilized more 
often than their relative abundance would dictate. Although the density of M. laticifera 
was greater in the open area (3.8 individuals/m?, SD = 1.4, n = 85) than at the forest edge 
(0.3 individuals/m2, SD = 1.2, n = 85), only 2 of the 150 plants sampled in the open area 
had early stages whereas 23 of the 60 plants sampled along the forest edge harbored eggs 
and/or larvae ( 7? = 56.4, df = 1, P = 0.001). 

Although little is known about the mechanisms of oviposition site selection in pierid 
butterflies, it is envisaged that females first locate appropriate habitats in which to search 
for host plants (Courtney 1986). Also, Pieridae frequently deposit eggs contagiously and 
may prefer plants growing at low densities (Root & Kareiva 1984, Courtney 1986). It 
seems that E. dina leuce females use the forest edge or some associated factor as a cue for 
finding host plants. The adaptative significance of this preference, if any, is unknown. 

Early instar larvae of E. dina leuce apparently avoid latex ducts by feeding between leaf 
veins of M. laticifera. Fourth and fifth instars observed in the field chewed a notch in the 
rachis and waited for the latex to drip from the wound prior to feeding on a leaf. Dussourd 


VOLUME 51, NUMBER 4 359 


(1993) suggested that caterpillars that feed on lactiferous plants sever veins and cut 
trenches specifically to deactivate the defensive function of pressurized latex. 

Although E. hecabe phoebus, E. elathea, and E. dina westwoodi use plants from families 
that characteristically produce latex (Jones et al. 1987, Brown 1992), the behavior of avoid- 
ing latex apparently has not been reported for Eurema. Moreover, Dussourd (1993), in a 
recent revision of caterpillar behavior for circumventing plant defenses, did not report any 
such case for pierids. This record for E. dina leuce suggests that complex behavior specifi- 
cally directed to circumvent latex defense may evolve relatively easily even in taxa appar- 
ently unassociated with lactiferous plants during much of their evolutionary histories. 

It is important to note that other legumes occurring in the study area [Chamaecrista 
patellaria Greene (Mimosaceae) and Stylosanthes viscosa SW (Fabaceae)]| are used as 
host plants by E. nise tenella Boisduval and E. elathea. Also E. albula Cramer feeds on four 
species of Senna sp. (Caesalpinaceae) at the same locality. Why E. dina leuce prefers to 
use a host plant that produces latex deserves further investigation. 

We conducted this study during the field course “Ecologia de Campo I” of the State 
University of Campinas (UNICAMP). K. S. Brown, Jr. identified the butterfly species, F. 
A. M. Santos made suggestions during the field work, and P. S. Oliveira, A. Shapiro and an 
anonymous reviewer provided helpful comments on the manuscript. Fellowship support 
to I. Andrade was provided by the Fundagao de Amparo a Pesquisa do Estado de Sao 
Paulo (grant number 95/2107-7). W. Benson was supported by a research fellowship from 
the CNPq. We thank the Companhia Vale do Rio Doce for essential logistic support and 
permission to work in the Linhares Forest Reserve. 


LITERATURE CITED 


BARNEBY, R. C. 1985. The identity and synonymy of Acacia guilandinae, Mimosa obo- 
vata, M. pseudo-obovata, and M. laticifera (Mimosaceae). Brittonia 37:85—87. 

BROWN, K. S. 1992. Borboletas da Serra do Japi: diversidade, habitats, recursos alimenta- 
res e variacao temporal, pp. 142-187. In L. P. C. Morellato (ed.), Histéria natural da 
Serra do Japi: ecologia e preservacgdo de uma area florestal no sudeste do Brasil. Edi- 
tora da UNICAMP / FAPESP. Campinas, Brazil. 

CLARK, G. C. & C. G. C. DICKSON. 1965. The life histories of two species of South 
African Eurema. J. Res. Lepid. 4:252—257. 

COURTNEY, S. P. 1986. The ecology of pierid butterflies: dynamics and interactions. Adv. 
Ecol. Res. 15:51-131. 

DEVRIES, P. J. 1985. Hostplant records and natural history notes on Costa Rican butter- 
flies (Papilionidae, Pieridae & Nymphalidae). J. Res. Lepid. 24:290—333. 

. 1987. The butterflies of Costa Rica and their natural history. Princeton Univ. 
Press, Princeton, New Jersey. 327 pp. 

Dussoub, D. E. 1993. Foraging with finesse: caterpillar adaptations for circumventing 
plant defenses, pp. 92-131. In N. E. Stamp & T. M. Casey (eds.), Caterpillars: ecolog- 
ical and evolutionary constraints on foraging. Chapman & Hall, New York. 

JONES, R. E. & J. RIENKS. 1987. Reproductive seasonality in the tropical genus Eurema 
(Lepidoptera: Pieridae). Biotropica 19:7—16. 

JONEs, R. E., J. RIENKS, L. WILSON, C. LOKKERS & T. CHURCHILL. 1987. Temperature, 
development and survival in monophagous and polyphagous tropical pierid butterflies. 
Austr. J. Zool. 35:235—246. 

Root, R. B. & P. M. KAREIvA. 1984. The search for resources by cabbage butterflies 
(Pieris rapae): ecological consequences and adaptive significance of Markovian move- 
ments in a patchy environment. Ecology 65:147—165. 

STANTON, M. L. 1982. Searching in a patchy environment: foodplant selection by Colias 
p. eriphyle butterflies. Ecology 63:839—853. 


ISABEL ANDRADE AND WOODRUFF W. BENSON, Graduate Program in Ecology, Univer- 
sidade Estadual de Campinas, 13083-970 Campinas, Sdo Paulo, Brazil 


Received for publication 6 August 1996; revised and accepted 2 December 1996. 


Journal of the Lepidopterists’ Society 
51(4), 1996, 360-363 


INDEX FOR VOLUME 51 


(new names in boldface) 


adamsi, Filatima, 32 brouana, Bagisara, 344 
Adamski, D., 32 Brown, John W., 193 
Aiello, Annette, 105 Brown, Richard L., 119 
Alibertia edulis, 105 Bucculatricidae, 227 
allozymes, 208 Bucculatrix 
americana, Genipa, 105 dominatrix, 227 
Anaea tetradymiae, 227 
eurypyle, 83 buckmoths, 47 
ryphea, 83 Buehner, Gretchen, 208 
Andaman Islands, 273 Burns, John, 1 
Ande, A. Taiwo, 269 
Andrade, Isabel, 358 Cadaret, Sean J., 208 
angelia, Electrostrymon, 184 Caldas, Astrid, 83 
Anisota senatoria, 208 California, 93, 119, 176, 227, 256, 333 
antennal color, 218 Callaghan, Curtis J., 57, 62 
Antheraea mylitta, 95, 187 Callophrys 
aposematism, 139, 149 eryphon, 176 
Arctiidae, 288 sheridanii, 75 
Arctostaphylos patula, 333 xami, 288 
Arizona, 197 Calydna volcanicus, 57 
arjuna, Terminalia, 95 cannibalism, 304 
artifical diet, 304 canusana, Phaneta, 119 
Aster ciliatus, 218 cardui, Vanessa, 256 
Asteraceae, 93, 227 Catling, Paul M., 218 
Asthenidia, 105 Catocala relicta, cover issue 2 
Austin, George T., 316 Celtis pallida, 197 
autumnal flight, 119 Cerastis 
azia, Ministrymon, 184 enigmatica, 227 
gloriosa, 227 
Baccharis pilularis, 227 robertsoni, 227 
bachmanii, Libytheana, 197 cervinaria, Synaxis, 333 
Bactra Chaetaglaea 
maiorana, 119 fergusoni, 135 
miwok, 119 sericea, 135 
priapeia, 119 tremula, 135 
Bagisara checklists, 272 
brouana, 344 Choristoneura fumiferana, 277 
praecelsa, 344 ciliatus, Aster, 218 
bait traps, 156 circadian rhythms, 75 
Baker, Crys, 288 Cirina forda, 269 
Balcazar L., Manuel A., 105 clinal variation, 47 
behavior, 288 clymene, Haploa, 288 
Benedetter, Andrea I., 208 Cnephasia longana, 93 
Benson, Woodruff W., 358 coastal chapparal, 227 
Berryman, Alan A., 333 collection donation, 102 
Betser, Cathleen L., 208 Colombia, 57, 62 
biogeography, 97, 176 Conner, William E., 288 
body weight, 91 conservation, 263 
book reviews, 191, 193 Cordero, Carlos, 288 
Borash, Jennifer L., 208 coremata, 288 
Bouhin, Herve, 75 courtship, 197, 288 
Brazil, 358 Crabo, Lars, 227 
Brou, Charlotte D., 159 crypsis, 333 


Brou, Vernon A., 135, 156 cyanira, Euselasia, 62 


VOLUME 51, NUMBER 4 


Davidson, Robert B., 288 
defoliator, 333 

dichotomous keys, 32, 62, 237 
dina, Eurema, 358 

dispersal, 176, 184, 208 
distribution, 156, 316 

diurnal species, 179 

diversity, 9 

dominatrix, Bucculatrix, 227 
Dryas iulia, 184 

Dryocampa rubicunda, cover issue 4 


eclosion, 75 

ecosystem dynamics, 263 
edulis, Alibertia, 105 

egg cannibalism, 304 

Eitan, Ofer, 197 
Electrostyrmon angelia, 184 
endangered species, 263, 273 
endemism, 97 
endocrinology, 75 
enigmatica, Cerastis, 227 
Erynnis, | 

eryphon, Callophrys, 176 
ethanol, 277 

Eurema dina, 358 

europa, Lethe, 273 
eurypyle, Anaea, 83 
Euselasia cyanira, 62 
evolution, 9 


Fasoranti, J. Olaniyan, 269 
faunal surveys/lists, 97, 249, 272, 316 
Ferguson, Douglas C., 344 
fergusoni, Chaetaglaea, 135 
fermentation, 277 
Ferrell, George T., 333 
Filatima 

adamsi, 32 

occidua, 32 

ornatifimbriella, 32 
fire, 263 
fitness, 277 
Florida, 184 
foodplants, 218, 269, 358 
forda, Cirina, 269 
Fountainea, 83 
French broom, 139, 149 
fumiferana, Choristoneura, 277 


Gelechiidae, 32 
gene-environment interaction, 95 
genetics, 208 

Genipa americana, 105 

Genista monspessulana, 139 
genitalia, 1, 32, 83, 316, 344 
geographic variation, 9 


Geometridae, 102, 333 
Gibson, Loran D., 119 
Giuliani, Derham, 256 
gloriosa, Cerastis, 227 
greenleaf manzanita, 333 
gregariousness, 208 
Grimble, David G., 97 
group living, 208 
Guatemala, 316 


hackberry, 197 
Hammond, Paul C., 97 
Haploa clymene, 288 
hawkmoths, 156, 91, 156 
Hemileuca 

lucina, 47 

maia, 47 

nevadensis, 47 
Hesperiidae, 1, 316 
Hiruma, Kiyoshi, 75 
Hodges, Ronald W., 32 
Homoeopteryx, 105 
host range, 139, 149 
hydroxydanaidal, 288 


imbibing, 277 

immature stages, 105 
Incisalia, 176 

incomptus, Niconiades, 316 
India, 95, 187, 273 
Inglorius, 316 

Inglorius mediocris, 316 
intoxication, 277 
intraspecific variation, 83 
introduced species, 119, 176 
iulia, Dryas, 184 


Japan, 304 
Johnson, Scott A., 208 


Karner Blue, 263 
Kelly, Scott D., 208 
Knebel, Andrea, 197 


Lafontaine, J. Donald, 227 
Lamiaceae, 93 

larval biology, 304, 358 
larval competition, 93 
larval ecology, 208 
leaf-miner, 227 

leaf-tier, 93 

lectotype, 32 

Leen, Rosemary, 139, 149 
legal protection, 273 
Leguminoseae, 358 

Lethe europa, 273 
Libytheana bachmanii, 197 


361 


362 


Libytheidae, 197 

life history, 105, 119, 269, 333 
light traps, 156 

longana, Cnephasia, 93 

Louisiana, 135, 159 

low temperature, 75 

lucina, Hemileuca, 47 

Lycaeides melissa, 263 
Lycaenidae, 75, 176, 184, 263, 288 


maia, Hemileuca, 47 
maiorana, Bactra, 119 
male competition, 288 
Malvaceae, 344 
management, 263 

mate locating behavior, 197 
mating behavior, 288 
McElveen, Melinda, 288 
mediocris, Inglorius, 316 
melissa, Lycaeides, 263 
Memphis, 83 

Mexico, 288 

Michigan, 47, 272 
migratory activity, 256 
Miller, William E., 9, 91, 277 
Mimosa, 358 

Ministrymon azia, 184 
Minnesota, 179 

miwok, Bactra, 119 
Mizohata, Hidetaka, 208 
modestia, Oxytenis, 105 
Mohanraj, Prasanth, 273 
monspessulana, Genista, 139 
Monterey pine, 176 


Museum of Comparative Zoology, 102 


mylitta, Antheraea, 95, 187 


nevadensis, Hemileuca, 47 
Nicobar Islands, 273 

Niconiades incomptus, 316 
Nigeria, 269 

Noctua pronuba, cover issue 3 
Noctuidae, 97, 102, 135, 237, 344 
non-target species, 97 
Nymphalidae, 83, 184, 218, 256 


oak barrens, 263 
occidua, Filatima, 32 
Ohio, 208 

Olwura, Takako, 304 
Ontario, 218 

Oregon, 97 
ornatifimbriella, Filatima, 32 
Osborne, Kendall H., 227 
oviposition, 218, 358 
Owens Valley, 256 
Oxytenis modestia, 105 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


pallida, Celtis, 197 
Panama, 105 

patula, Arctostaphylos, 333 
Pelham, Jonathan P., 75 
Perkins, Owen A., 272 
Phaneta canusana, 119 
phenology, 156, 218, 269, 333 
pheromones, 249 
Phyciodes tharos, 218 
phylogeny, 9, 83, 105 
Pierce, Naomi, 102 
Pieridae, 304, 358 

Pieris rapae, 304 

pilularis, Baccharis, 227 
plant latex, 358 

pollination, 9 

population dynamics, 256 
Porter, Adam H., 208 
Powell, Jerry A., 93, 176 
praecelsa, Bagisara, 344 
prairies, 179 

prescribed burning, 263 
priapeia, Bactra, 119 
pronuba, Noctua, cover issue 3 
pupal diapause, 75 
Pyralidae, 139, 149 
pyrrolizidine alkaloids, 288 


quinolizidine alkaloids, 139, 149 


range expansion, 119 

rapae, Pieris, 304 

Reed, Catherine C., 179 
relatedness, 208 

relicta, Catocala, cover issue 2 
reproductive adaptation, 187 
reversalis, Uresiphita, 149 
Riodinidae, 57, 62 
robertsoni, Cerastis, 227 
Rubiaceae, 105 


rubicunda, Dryocampa, cover issue 4 


Rubinoff, Daniel Z., 227 
Rutowski, Ronald L., 197 
ryphea, Anaea, 83 


Salazar, Julian, 57 
Santa Rosa Island, 93 


Saturniidae, 47, 95, 105, 187, 208, 269 


Satyridae, 273 

Scholtens, Brian G., 47 
Scrophulariaceae, 93 
senatoria, Anisota, 208 
Sengupta, A. K., 95 
sericea, Chaetaglaea, 135 
Sesiidae, 249 

sheridanii, Callophrys, 75 
Sherman, Marilyn L., 208 


VOLUME 51, NUMBER 4 


Shields, Oakley, 256 
Shuey, John A., 191, 263 
Siddiqui, A. A., 95 

silk yield, 95 

species richness, 179 
sphagnum bogs, 227 
Sphingidae, 9, 91, 156 
starvation, 304 

status assessment, 273 
surveys, 179 

Synaxis cervinaria, 333 


tameamea, Vanessa, cover issue | 
tasar silk moth, 95, 187 


taxonomy, 32, 47, 57, 62, 83, 119, 135, 227, 


237, 316, 344 
Terkanian, Barbara, 197 
Terminalia arjuna, 95 
territoriality, 288 
tetradymiae, Bucculatrix, 227 
tharos, Phyciodes, 218 
tongue length, 9 
Tortricidae, 93, 119, 128, 277 
tremula, Chaetaglaea, 135 


Uresiphita, 139, 149 
Uresiphita reversalis, 149 


vagility, 179 
Valenti, Michael A., 333 
Vanessa 
cardui, 256 — 
tameamea, cover issue | 
Veenakumari, K., 273 
vernal flight, 119 
Vinpeius, 316 
volcanicus, Calydna, 57 
voltinism, 135, 156 


Wagner, Warren H., Jr., 47 
Washington, 75 

Watanabe, Mamoru, 304 
weather, 256 

West Virginia, 249 

wing length, 91 

wing pattern, 83 

winter moths, 135 

Winter, William D., 102 
Wright, Donald J., 119 


xami, Callophrys, 288 
Xanthodes, 344 


Date of Issue (Vol. 51, No. 4): 29 December 1997 


363 


EDITORIAL STAFF OF THE JOURNAL 


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Peabody Museum of Natural History 
Yale University 
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email: lawrence.gall@yale.edu 


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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


INTOXICATED LEPIDOPTERANS: HOW IS THEIR FITNESS AFFECTED, AND WHY DO 
THEY TIPPLE? . William E. Miller. ee ae 


HypROXYDANAIDAL AND THE COURTSHIP OF HapLoa (ArcTiuDAE) Robert B. 


Davidson, Crys Baker, Melinda McElveen and William E. Conner _. 


Wuy DO SOME MALE GALLopHRYS XAMI (LYCAENIDAE) SHIFT THEIR TERRI- 
ToRtES?. « Carlos Gorderdic 2 es 


EGG CANNIBALISM BY NEWLY HATCHED LARVAE OF THE SMALL WHITE BUTTER- 
FLY, PIERIS RAPAE CRUCIVORA (PIERIDAE), ON AN ARTIFICIAL DIET Mamoru 


Watanabe and Takako Ohiurd 0 A ee 


NoTeEs ON HESPERIIDAE IN NORTHERN GUATEMALA, WITH DESCRIPTIONS OF NEW 
TAXA George T. Austin 6008 i eS 


LIFE HISTORY AND BEHAVIOR OF SYNAXIS CERVINARIA (GEOMETRIDAE), A DEFOLIATOR 
oF ARCTOSTAPHYLOS PATULA (ERICACEAE) Michael A. Valenti, Alan A. 
Berryman and George T. Ferrell. 0 2 SS ee 


REVIEW OF THE NEw Wokr.p BaAGISARINAE WITH DESCRIPTION .OF TWO NEW 
SPECIES FROM THE SOUTHERN Unitep States (Noctumpae) -Douglas C. 
OV OUSON a a0 Nl: Es Wek Oe oe ON ae Ne eiet e 


GENERAL NOTES 


Eurema dina leuce (Pieridae) feeds on a lactiferous hostplant in eastern Brazil 
Isabel Andrade and Woodruff W. Benson.22 2 


INDEX: FOR. VOLUME O12 22 0 


288 


295 


304 


316 


333 


344 


This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). - 


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